End of season carbon supply status of woody species near the treeline in western China

As trees and shrubs approach the high elevation tree limit, it is often assumed that they fall short in photosynthate (source limitation). Alternatively, low temperature may restrict carbon investment (growth, sink limitation). The content of mobile non-structural carbohydrates (NSC) in tissues is considered a measure of the carbon source–sink balance. To test the source vs. sink limitation hypothesis, we compared late-season NSC concentrations of various woody taxa across altitudinal gradients from the subalpine forest to the treeline at the eastern edge of the Tibetan Plateau. Since we were interested in the generality of trends, we present “community” trends across four taxa, namely Quercus aquifolioides, Abies faxoniana, Rhododendron fabri subsp. prattii and Sorbus rufopilosa. NSC concentrations increased significantly with altitude in branch wood, current-year and last-year leaves, while there were no significant trends in stem sapwood and root xylem. The sugar to starch ratio was roughly 1:1 in branches and evergreen leaves, while stems and roots showed a higher starch fraction. Analyses of total nitrogen in leaves and wood tissues indicated no change in the trees’ nitrogen supply with elevation. The overall altitudinal trends of NSC in this group of woody plant species revealed no depletion of carbon reserves near the tree limit, suggesting that sink limitation predominates woody plant life across this treeline ecotone community.     

秦岭东段不同海拔栓皮栎粗根非结构性碳水化合物含量的季节动态

掌握树木根部碳存储规律对于准确估算碳在地上器官与地下器官间的分配非常必要。本研究以栓皮栎(Quercus variabilis Blume)为对象,在2016年5月—2017年6月,通过周期性采样方法(共计采样14次),测定了高、低海拔(970和650 m)栓皮栎粗根非结构性碳水化合物(non-structural carbohydrates,NSC)及其组分(可溶性糖和淀粉)含量的年内动态变化。结果表明:除高海拔淀粉外,栓皮栎粗根NSC及其组分含量均随季节变化差异显著(P<0.05)。粗根NSC含量呈现生长季初期(3月)下降,非生长季(2月)达到最高值的变化趋势;栓皮栎粗根NSC组成以淀粉为主,高、低海拔淀粉含量占比分别为61%和71%,这可能与栓皮栎适应区域环境特征有关。不同海拔间,栓皮栎粗根NSC及其组分含量的差异出现在生长季初期(3月,P<0.05)。高海拔(10.26%)栓皮栎粗根NSC含量小于低海拔(13.96%)。栓皮栎粗根NSC含量存在明显的季节波动,粗根在生长季末及非生长季积累的NSC对下一年树木生长启动非常重要,研究结果有助于理解树木地下器官对树木生长的碳供应机制。     

Carbon dynamics in the deciduous broadleaf tree Erman's birch (Betula ermanii) at the subalpine treeline on Changbai Mountain,Northeast China

Premise of the Study

The growth limitation hypothesis (GLH) and carbon limitation hypothesis (CLH) are two dominant explanations for treeline formation. The GLH proposes that low temperature drives the treeline through constraining C sinks more than C sources, and it predicts that non‐structural carbohydrate (NSC) levels are static or increase with elevation. Although the GLH has received strong support globally for evergreen treelines, there is still no consensus for deciduous treelines, which experience great asynchrony between supply and demand throughout the year.

Methods

We investigated growth and the growing‐season C dynamics in a common deciduous species, Erman's birch (Betula ermanii), along an elevational gradient from the closed forest to the treeline on Changbai Mountain, Northeast China. Samples were collected from developing organs (leaves and twigs) and main storage organs (stems and roots) for NSC analysis.

Key Results

Tree growth decreased with increasing elevation, and NSC concentrations differed significantly among elevations, organs, and sampling times. In particular, NSC levels varied slightly during the growing season in leaves, peaked in the middle of the growing season in twigs and stems, and increased continuously throughout the growing season in roots. NSCs also tended to increase or vary slightly in developing organs but decreased significantly in mature organs with increasing elevation.

Conclusions

The decrease in NSCs with elevation in main storage organs indicates support for the CLH, while the increasing or static trends in new developing organs indicate support for the GLH. Our results suggest that the growth limitation theory may be less applicable to deciduous species' growth than to that of evergreen species.     

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.     

Patterns in nonstructural carbohydrate contents at the tree organ level in response to drought duration

Nonstructural carbohydrates (NSCs) facilitate the adaptation of trees to drought stress. There have been a large number of studies exploring NSC changes in individual plant species and individual organ under drought and showed different trends; however, an understanding of the universal pattern of the plant NSCs responses to drought, particularly to drought duration, is still lacking. Here, we compiled data from 47 experimental studies on 52 tree species and conducted a meta‐analysis to evaluate the responses of soluble sugars, starch, and TNSC (total nonstructural carbohydrates including both soluble sugars and starch) concentrations in different tree organs (leaf, stem, and root) to drought intensity and duration. We found that starch in all organs decreased and soluble sugars in leaf increased with prolonged experiment time, and the changes in soluble sugars in all organs were stronger under severe drought than under slight‐to‐moderate drought. Under slight‐to‐moderate drought, the NSC content of each organ varied with time, while with the extension of the drought duration, the NSCs gradually approached the control value (no drought stress); this trend remained in the late drought, which means that trees activated physiological regulation processes to increase carbon storage and reduce the risks of carbon starvation. In contrast, long‐term severe drought could lead to a net loss of carbohydrates, especially in the root, implying that prolonged severe drought could lead to NSC depletion in the whole plant. As prolonged drought duration has occurred in and is projected for many regions, this paper could shed light into studies on how trees respond and adapt extending drought duration through nonstructural carbon production, transportation, and reallocation.     

Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients

Background and Aims

The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous–evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations.

Methods

Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous–evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra).

Key Results

Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood.

Conclusions

Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone.     

No evidence for depletion of carbohydrate pools in Scots pine (Pinus sylvestris L.) under drought stress

The physiological mechanisms leading to Scots pine (Pinus sylvestris L.) decline in the dry inner alpine valleys are still unknown. Testing the carbon starvation hypothesis, we analysed the seasonal course of mobile carbohydrate pools (NSC) of Scots pine growing at a xeric and a dry-mesic site within an inner alpine dry valley (750 m a.s.l., Tyrol, Austria) during 2009, which was characterised by exceptional soil dryness. Although, soil moisture content dropped to ca. 10% at both sites during the growing season, NSC concentrations rose in all tissues (branch, stem, root) until the end of July, except in needles, where maxima were reached around bud break. NSC concentrations were not significantly different in the analysed tissues at the xeric and the dry-mesic site. At the dry-mesic site, NSC concentrations in the aboveground tree biomass were significantly higher during the period of radial growth. An accumulation of NSC in roots at the end of July indicates a change in carbon allocation after an early cessation in aboveground growth, possibly due to elevated belowground carbon demand. In conclusion, our results revealed that extensive soil dryness during the growing season did not lead to carbon depletion. However, even though carbon reserves were not exhausted, sequestration of carbohydrate pools during drought periods might lead to deficits in carbon supply that weaken tree vigour and drive tree mortality.     

海拔梯度对祁连山青海云杉林乔木层和土壤层碳密度的影响

以祁连山西水林区青海云杉典型林分为研究对象,按照青海云杉分布界限海拔2500—3300 m,采用梯度格局法,研究祁连山青海云杉林乔木层和土壤层碳密度沿海拔梯度的空间分布特征,以期为准确估算祁连山青海云杉林碳储量变化影响因素提供科学依据。结果表明:(1)青海云杉林生物量平均值为115.83 t/hm~2,碳密度平均值为60.23 t/hm~2。生物量整体随海拔梯度增加表现为先增加后波动降低的趋势,在海拔2800 m处达到最高值(197.10 t/hm~2),海拔3300 m处达到最低值(7.66t/hm~2),且不同海拔梯度间差异显著。林分各器官生物量分配格局在各海拔处均表现为干根枝叶。(2)土壤有机碳含量平均值为54.80 g/kg,变化范围为31.49—76.96 g/kg。随着土壤层次的增加,除海拔3200 m和3300 m的土壤有机碳含量未表现出规律变化外,其他海拔梯度则均呈现出逐渐降低趋势。土壤有机碳密度在海拔2900 m最高,为245.40 t/hm~2,在海拔2700 m处最低,为130.24 t/hm~2;海拔2500—2700 m表现为平缓降低趋势,在2800 m处急剧上升,且海拔2800—3200 m呈现无显著性轻度波动变化,在海拔3300 m又急剧降低。(3)青海云杉林生态系统平均总碳密度为255.15 t/hm~2,乔木层和土壤层占总碳密度的比例分别为23.61%和76.39%,且不同海拔梯度间存在极显著差异。土壤有机碳密度与海拔、年均降水量、土壤有机碳含量、土壤全氮呈显著正相关,与年夏季平均气温呈显著负相关;乔木层碳密度与年夏季气温、林分密度、胸高断面积呈显著正相关,与海拔和土壤全氮呈显著负相关。(4)祁连山青海云杉林乔木层和土壤层碳密度均随海拔梯度变化受水热条件组合的改变而呈现规律变化,以中部海拔区段2800—3200 m碳密度较高。     

Effect of size and altitude on soil organic carbon stock in homegarden agroforestry system in Central Himalaya,India

Homegarden agroforestry system has potential to sequester carbon while simultaneously producing agriculture yield and tree products. This study analyzed relationship between homegarden size and soil organic carbon stock (SCS) along an altitudinal gradient in Central Himalaya, India. Homegardens were selected at four altitudes (very low, low, mid and high) and categorized into three size classes viz. small, medium and large. Soil samples from surface (0–15 cm) and sub surface soils (15–30 cm) were collected seasonally and analyzed. Results indicated that SCS ranged between 22.56 and 81.51 t C ha⁻¹ and increased with increasing altitude for small homegardens. For medium and large homegardens, it increased up to mid altitude and then decreased in high altitude homegardens. At each altitude, maximum SCS (38.56–66.16 t C ha⁻¹) was recorded for medium sized homegardens except at high altitude where it was maximum (66.01 t C ha⁻¹) for small homegardens. Seasonal variation in SCS showed inconsistent pattern along the altitudinal gradient. The differences in SCS due to size and altitude recorded in the present study are possibly caused by distinct quality of plant material and management practices.     

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.     

Hydraulics play an important role in causing low growth rate and dieback of aging Pinus sylvestris var. mongolica trees in plantations of Northeast China

The frequently observed forest decline in water‐limited regions may be associated with impaired tree hydraulics, but the precise physiological mechanisms remain poorly understood. We compared hydraulic architecture of Mongolian pine (Pinus sylvestris var. mongolica) trees of different size classes from a plantation and a natural forest site to test whether greater hydraulic limitation with increasing size plays an important role in tree decline observed in the more water‐limited plantation site. We found that trees from plantations overall showed significantly lower stem hydraulic efficiency. More importantly, plantation‐grown trees showed significant declines in stem hydraulic conductivity and hydraulic safety margins as well as syndromes of stronger drought stress with increasing size, whereas no such trends were observed at the natural forest site. Most notably, the leaf to sapwood area ratio (LA/SA) showed a strong linear decline with increasing tree size at the plantation site. Although compensatory adjustments in LA/SA may mitigate the effect of increased water stress in larger trees, they may result in greater risk of carbon imbalance, eventually limiting tree growth at the plantation site. Our results provide a potential mechanistic explanation for the widespread decline of Mongolian pine trees in plantations of Northern China.     

芦芽山不同海拔白杄非结构性碳水化合物含量动态

高山林线对环境变化具有高度的敏感性, 但林线形成机制仍然没有明确的结论。为了检验高山林线形成是由碳限制还是生长限制决定, 并探讨林线树种适应高山环境的生理生态机制, 选择山西省吕梁山脉北端芦芽山, 沿3个海拔梯度测定了林线树种白杄(Picea meyeri)各组织非结构性碳水化合物(NSC)及其组分含量。结果表明: 白杄总体及各组织NSC含量均随海拔升高而增加, 林线树木不存在碳限制; 白杄NSC源、汇均随海拔升高而增加, 源-汇比在3个海拔之间没有差异, 表明源-汇平衡关系对海拔的适应性, 林线树木碳源活动没有受到限制; 各组织中可溶性糖与淀粉的比值随海拔升高呈增大趋势, 说明树木生长的环境越寒冷, 树木组织中表现出越明显的保护策略, 也可能暗示林线区域的树木更多地受到生长限制。研究结果在一定程度上支持“生长限制”假说。     

Growth decline of Pinus Massoniana in response to warming induced drought and increasing intrinsic water use efficiency in humid subtropical China

Pinus Massoniana is the most widely distributed coniferous species in southern China and one of the most distributed species for plantation in China. It is not uncertain about the responses of tree growth to the combined effects of regional drying and the increase in the intrinsic water-use efficiency (iWUE) due to increased atmospheric CO₂. This study addressed this issue by comparing the tree growth patterns as represented by three tree-ring width chronologies with climate variables and three iWUE series derived from three tree-ring stable carbon isotope discrimination chronologies (Δ¹³C) from Pinus Massoniana in Daiyun Mountain, central Fujian province of China. Among these chronologies, we reported the first tree-ring carbon isotope discrimination chronologies (Δ¹³C) from Fuzhou area spanning last 210 years. It was found that tree radial growth is mainly limited by dry condition from May to October. Growth limitation by cold condition was only found in one high altitude site (780m) in early spring and late autumn. The tree-ring carbon discrimination was enhanced under conditions with low relative humidity and sufficient sunshine in late summer and autumn. In general, the iWUE showed a significantly increasing trend since the 1850s for all the sites in response to the increase in atmospheric CO₂. However, the growth promotion of the increased iWUE on tree growth could not compensate the growth limitation caused by drought. Especially since the 1960s, a growth decline was found at two drought stressed sites at low altitudes. On the other hand, the increase in temperature of spring and autumn and iWUE has most likely enhanced tree growth at the high altitude site.     

青藏高原东南缘陇蜀杜鹃当年生小枝茎叶性状对海拔变化的响应

通过在青藏高原东南缘的岷江源区三处高山林草交错带采集陇蜀杜鹃当年生小枝,测量小枝的茎长、茎粗与茎生物量,沿海拔梯度自然分布的空间变化对比其当年生小枝的茎结构与生物量及其相互关系,接着比较了总叶质量和比叶面积的变化,最后通过典范对应分析(CCA)探究小枝茎叶性状与土壤因子的相关关系,有助于理解陇蜀杜鹃适应异质环境的生态适应对策。结果表明:(1)除弓杠岭外,斗鸡台与卡卡山土壤因子在海拔间差异显著,相似海拔不同山地间土壤因子也存在显著差异。(2)除卡卡山高海拔比叶面积显著小于低海拔外,其他样地的小枝茎性状(茎长、茎粗、比茎长和茎生物量)和叶性状(总叶质量和比叶面积)在海拔间并无显著差异,相似海拔不同坡体间茎粗、茎生物量以及比茎长差异也未达到显著水平(P>0.05),但弓杠岭(3963 m)的茎长显著大于卡卡山(3921 m)(P<0.05)。(3)小枝茎长、茎粗与茎生物量之间均为异速生长关系,且茎生物量累积速率大于茎长和茎粗增加的速率,且茎长的增加速率约为茎粗的2.8倍。(4)CCA排序结果表明,11个土壤环境因子中,碳含量、碳氮比、有机碳含量、总磷和容重对植物性状有显著的影响,但在海拔及不同山体间影响小枝茎叶性状的土壤因子不同。由此可知,较小的海拔梯度变化对小枝茎叶性状影响较小,土壤因子是导致小枝茎叶性状差异的重要原因之一。     

Effects of increasing root carbon investment on the mortality and resprouting of Haloxylon ammodendron seedlings under drought

• Tree mortality induced by drought is one of the most complex processes in ecology. Although two mechanisms associated with water and carbon balance are proposed to explain tree mortality, outstanding problems still exist.
• Here, in order to test how the root system benefits survival and resprouting of Haloxylon ammodendron seedlings, we examined the various water‐ and carbon‐related physiological indicators (shoot water potential, photosynthesis, dark respiration, hydraulic conductance and non‐structural carbohydrates [NSC]) of H. ammodendron seedlings, which were grown in drought and control conditions throughout a grow season in greenhouse.
• The survival time of the seedling root system (died 70 days after drought) doubled the survival time of the shoot (died at 35 days). Difference in survival time between shoot and root resulted from sustained root respiration supported by increased NSC in roots under drought. Furthermore, investment into the root contributed to resprouting following drought. Based on these results, a death criterion is proposed for this species. The time sequence of major events indicated that drought shifted carbon allocation between shoot and root and altered the flux among different sinks (growth, respiration or storage). The interaction of water and carbon processes determined death or survival of droughted H. ammodendron seedlings.
• These findings revealed that the ‘root protection’ strategy is critical in determining survival and resprouting of this species, and provided insights into the effects of carbon and water dynamics on tree mortality.

     

Disturbance from traditional fire management in subalpine heathlands increases Afro‐alpine plant resilience to climate change

Species are often controlled by biotic factors such as competition at the warm edge of their distribution range. Disturbances at the treeline, disrupting competitive dominance, may thus enable alpine species to utilize lower altitudes. We searched for evidence for range expansion in grazed, fire‐managed Ethiopian subalpine Erica heathlands across a 25‐year chronosequence. We examined vascular plant composition in 48 plots (5 × 5 m) across an altitudinal range of 3,465–3,711 m.a.s.l. and analyzed how community composition changed in relation to increasing competition over time (using a Shade index based on Erica shrub height and cover) and altitude. Species‘ habitats and altitudinal ranges were derived from literature. Time since fire explained more variation (r² = .41) in species composition than altitude did (r² = .32) in an NMDS analysis. Community‐weighted altitudinal optima for species in a plot decreased strongly with increasing shade (GLM, Standardized Regression Coefficient SRC = ?.41, p = .003), but increased only weakly with altitude (SRC = .26, p = .054). In other words, young stands were dominated by species with higher altitudinal optima than old stands. Forest species richness increased with Log Shade index (SRC = .12, p = .008), but was unaffected by altitude (SRC = ?.07, p = .13). However, richness of alpine and heathland species was not highest in plots with lowest Shade index, but displayed a unimodal pattern with an initial increase, followed by a decrease when shading increased (altitude was not significant). Our results indicate that disturbance from the traditional patch burning increases the available habitat for less competitive high‐altitude plants and prevents tree line ascent. Therefore, maintaining, but regulating, the traditional land use increases the Afro‐alpine flora's resilience to global warming. However, this system is threatened by a new REDD+ program attempting to increase carbon storage via fire suppression. This study highlights the importance of understanding traditional management regimes for biodiversity conservation in cultural landscapes in an era of global change.     

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.     

Effects of altitude and livestock on the regeneration of two tree line forming <Emphasis Type="Italic">Polylepis</Emphasis> species in Ecuador

Regeneration is known to be limited at many temperate tree lines, but very little data is available on the impacts of altitude and anthropogenic disturbance on regeneration patterns along tropical tree lines. The study focused on the reproductive traits of two Polylepis species in the Páramo de Papallacta in Ecuador along an altitudinal gradient, and involved different intensities of cattle trampling within subsequent altitudinal ranges. We analyzed flowering, fruit set, seed viability, germination, and seedling establishment as well as stand structure of Polylepis incana and P. pauta. The numbers of P. incana inflorescences and seedlings per m2 showed a marginally significant decrease with increasing altitude. Mean tree height was significantly lower at higher altitudes, while stem number increased. The number of P. pauta inflorescences also decreased significantly upslope. In both forest types, trampling was found to have a positive impact on seedling abundance, presumably due to the removal of the litter layer. Thus, there was no evidence of negative effects of moderate cattle grazing on both tree line species. However, sapling establishment was minimal inside the forest stands at all altitudes and grazing levels, and we consequently observed a low proportion of narrow stems within all investigated forests. Our results show that, along with vegetative growth limitations of adult trees, important regeneration traits such as seedling and inflorescence numbers are also influenced by altitude, which might contribute to the formation of the upper tree line. Nevertheless, recruitment in the forest interior was low overall indicating that further factors, such as light conditions, affect regeneration of the studied species.     

Non-structural carbon compounds in temperate forest trees

The current carbon supply status of temperate forest trees was assessed by analysing the seasonal variation of non‐structural carbohydrate (NSC) concentrations in leaves, branch wood and stem sapwood of 10 tree species (six deciduous broad‐leafed, one deciduous conifer and three evergreen conifer trees) in a temperate forest that is approximately 100 years old. In addition, all woody tissue was analysed for lipids (acylglycerols). The major NSC fractions were starch, sucrose, glucose and fructose, with other carbohydrates (e.g. raffinose and stachyose) and sugar alcohols (cyclitols and sorbitol) playing only a minor quantitative role. The radial distribution of NSC within entire stem cores, assessed here for the first time in a direct interspecific comparison, revealed large differences in the size of the active sapwood fraction among the species, reflecting the specific wood anatomy (ring‐porous versus diffuse‐porous xylem). The mean minimum NSC concentrations in branch wood during the growing season was 55% of maximum, and even high NSC concentrations were maintained during times of extensive fruit production in masting Fagus sylvestris. The NSC in stem sapwood varied very little throughout the season (cross species mean never below 67% of maximum), and the small reductions observed were not significant for any of the investigated species. Although some species contained substantial quantities of lipids in woody tissues (‘fat trees’; Tilia, Pinus, Picea, Larix), the lipid pools did not vary significantly across the growing season in any species. On average, the carbon stores of deciduous trees would permit to replace the whole leave canopy four times. These data imply that there is not a lot of leeway for a further stimulation of growth by ongoing atmospheric CO₂ enrichment. The classical view that deciduous trees rely more on C‐reserves than evergreen trees, seems unwarranted or has lost its justification due to the greater than 30% increase in atmospheric CO₂ concentrations over the last 150 years.     

Restoring for the present or restoring for the future: enhanced performance of two sympatric oaks (Quercus ilex and Quercus pyrenaica) above the current forest limit

Reforestation is common to restore degraded ecosystems, but tree‐species choice often neglects ongoing environmental changes. We evaluated the performance of planted seedlings of two oak species at two sites in a Mediterranean mountain (Sierra Nevada, SE Spain): one located within the current altitudinal forest range (1,600–1,760 m), and one above the upper forest limit (1,970–2,120 m). The forest service planted 1,350 seedlings of the deciduous Pyrenean oak and the evergreen Holm oak in a postfire successional shrubland. After 2 years, seedlings were monitored for survival, and a subset of 110 Pyrenean oaks and 185 Holm oaks were harvested for analyses of biomass and foliar nutrient status, δ¹³C, and δ¹⁸O. Both species showed the highest survival and leaf N status above the upper forest limit, and survival increased with altitude within each plot. The deciduous oak benefited most from planting at higher altitude, and it also had greater biomass at the higher site. Correlations between foliar N, δ¹⁸O, and δ¹³C across elevations indicate tighter stomatal control of water loss and greater water‐use efficiency with increasing plant N status at higher altitude, which may represent a so‐far overlooked positive feedback mechanism that could foster uphill range shifts in water‐limited mountain regions. Given ongoing trends and future projections of increasing temperature and aridity throughout the Mediterranean region, tree‐species selection for forest restoration should target forecasted climatic conditions rather than those prevailing in the past. This study highlights that ecosystem restoration provides an opportunity to assist species range shifts under rapidly changing climate.