高山林线变化的更新受限机制研究进展

全球林线位置对气候变暖的响应表现为上升、无变化或下降等截然不同趋势,表明影响林线位置及动态的因子十分复杂,除了较普遍认为的低温调控机制外,还存在其它控制林线位置变化的机制。林线向上迁移开始于种子向林线以上的传播及幼苗在林线以上的定居,这些过程中的限制因子均会影响林线的位移,因此研究更新过程及其限制因子对理解高山林线对气候变化的响应具有重要的科学意义。主要从种子和幼苗两个关键阶段综述高山林线森林更新的研究进展。在种子阶段,夏季积温不足导致种子产量和活力下降,风速过低和浓密灌丛限制种子向林线以上传播,近地表的霜冻/水分胁迫和灌木释放的化感物质会阻碍种子在林线以上萌发。在幼苗阶段,除冬季低温外,生长季内较大的温度日振幅和偶然出现的冻害事件也是导致幼苗死亡的重要原因,而低温环境下的强烈光照引起的低温光抑制会显著降低生长季的光合作用;土壤低温、由土壤温度昼夜变化引起的冻举事件、夏季土壤干旱可能会导致幼苗光合作用下降和死亡率上升;积雪太浅会导致生长季早期幼苗水分供应的严重缺乏,但积雪太深会导致幼苗感染真菌的可能性增加;浓密的灌木和草本植物以及植食动物的啃食也会降低林线以上的幼苗存活率。气候变暖对林线幼苗定居的影响复杂且具有很大不确定性,需要进一步研究气候变暖导致的环境因子变化对林线更新各关键阶段的影响。未来气候变暖无疑会导致生长季起始日提前,结束日推迟,这很可能会增加生长季期间尤其是早期的低温冻害事件,对高山林线树种幼苗的存活具有重要影响。在未来研究中,需要找出定义生长季冻害事件的温度阈值,利用长期气象观测数据分析增温背景下生长季早期冻害事件特征的变化趋势,并进一步开展野外模拟增温实验以深刻理解林线树种的种子萌发和幼苗定居与生长季冻害事件的关系,加强对不同地区林线树种的繁殖策略研究,这将有助于人们进一步理解不同区域林线的形成机制并预测未来气候变化条件下林线的动态变化趋势。     

高山林线生态交错区木本植物幼苗分布特征、更新机制及其对气候变化的响应

木本植物幼苗是高山林线生态交错区的重要组成部分,其更新对气候变化背景下树线的移动至关重要.本研究通过对近几十年来全球范围内林线生态交错区的木本植物幼苗分布特征、更新机制及其对气候变化响应的研究总结得出:林线生态交错区木本植物幼苗的空间分布类型主要为渐变型和聚集型,且不同分布类型对树线动态的指示意义各异.在全球尺度上,其分布的海拔高限通常与生长季长度、均温和物种特性等有关,而在区域尺度上则多受降水影响.在幼苗更新初期,种源在很大程度上决定了种子的萌发及分布位置,之后微环境的促进作用为幼苗的定植提供庇护,提高其存活率,而在更新后期多种生物和非生物因素及其相互作用则非常关键.气候变暖促使林线生态交错区气温升高、降水充沛,有利于幼苗生长,使其向高海拔区域扩张而成为树线上移的先兆,但部分物种受遗传特性或适应策略影响,仅表现为密度增加,使树线保持相对稳定.未来应借助树轮、14C等精确定年技术,通过长期的野外定位观测和室内模拟,加强多时空尺度下林线幼苗的空间分布特征和更新机制研究,分析不同类型林线内木本植物幼苗的适应策略,预测气候变化背景下的树线动态,为山地生态系统恢复及保护提供科学依据.     

贡嘎山雅家埂峨眉冷杉林线种群的时空动态

通过对贡嘎山雅家埂峨眉冷杉种群林线附近6个3000 m2样地(阴阳坡各3个)中峨眉冷杉(Abies fabri Craib)种群的定位调查,分析了过去100a间该区峨眉冷杉种群的时间-空间动态。结果表明:1)雅家埂林线附近峨眉冷杉种群密度在过去100 a(主要是近50 a)有显著的升高,但树线的海拔位置并无明显的爬升;2)阴阳坡林线格局存在显著的坡向分异:阴坡林线和树线的海拔高度显著高于阳坡(分别比阳坡高152.5 m和135.8 m),阳坡林线附近峨眉冷杉早期的生长速率在大于阴坡,但后期的生长速率却低于阴坡;3)热量(温度)控制假说不能完全解释雅家埂目前的树线格局,除气候因素之外,其它因素也限制了雅家梗地区树线位置的变化。     

Gradual expansion of moisture sensitive Abies spectabilis forest in the Trans-Himalayan zone of central Nepal associated with climate change

Population structure and tree recruitment dynamics in the natural treeline ecotone of high mountains are strong indicators of vegetation responses to climate. Here, we examined recruitment dynamics of Abies spectabilis across the treeline ecotone (3439–3638 m asl) of Chimang Lekh of Annapurna Conservation Area in the Trans-Himalayan zone of central Nepal. Dendrochronological techniques were used to establish stand age structure by ring counts of adults, and by terminal bud scar count for seedlings and saplings. The results showed abundant seedling recruitment, higher regenerative inertia and colonization with a consistent range shift of the A. spectabilis treeline. The upward expansion of this sub-alpine treeline was found to be driven by a strong dependence of seedling recruitment and radial growth on snowmelt and precipitation as temperatures rise. The radial growth of A. spectabilis at the alpine timberline ecotone (ATE) and closed timberline forest (CTF) showed sensitivity to spring season (March–May) climate. Tree ring indices of CTF showed a strong positive correlation with spring and annual precipitation, and a significant negative correlation with spring and annual temperature, however, moisture sensitivity was less strong at ATE than CTF.     

Temporal dynamic of wood formation in Pinus cembra along the alpine treeline ecotone and the effect of climate variables

We determined the temporal dynamic of cambial activity and xylem development of stone pine (Pinus cembra L.) throughout the treeline ecotone. Repeated micro-sampling of the developing tree ring was carried out during the growing seasons 2006 and 2007 at the timberline (1,950 m a.s.l.), treeline (2,110 m a.s.l.) and within the krummholz belt (2,180 m a.s.l.) and the influence of climate variables on intra-annual wood formation was determined. At the beginning of both growing seasons, highest numbers of cambial and enlarging cells were observed at the treeline. Soil temperatures at time of initiation of cambial activity were c. 1.5°C higher at treeline (open canopy) compared to timberline (closed canopy), suggesting that a threshold root-zone temperature is involved in triggering onset of above ground stem growth. The rate of xylem cell production determined in two weekly intervals during June through August 2006–2007 was significantly correlated with air temperature (temperature sums expressed as degree-days and mean daily maximum temperature) at the timberline only. Lack of significant relationships between tracheid production and temperature variables at the treeline and within the krummholz belt support past dendroclimatological studies that more extreme environmental conditions (e.g., wind exposure, frost desiccation, late frost) increasingly control tree growth above timberline. Results of this study revealed that spatial and temporal (i.e., year-to-year) variability in timing and dynamic of wood formation of P. cembra is strongly influenced by local site factors within the treeline ecotone and the dynamics of seasonal temperature variation, respectively.     

Comparison of growth-climate relationship of sabina przewalskii at different timberlines along a precipitation gradient in the northeast Qinghai-Xizang Plateau,China北大核心CSCD

Aims To test the hypothesis that water is the main limiting factor of tree growth at the arid alpine timberline, and to explore the effects of water on growth-climate relationships of Sabina przewalskii along a precipitation gradient in the northeast Qinghai-Xizang Plateau. Methods Three sides were selected to sample the alpine timberline along a precipitation gradient in the northeast Qinghai-Xizang Plateau: Halihatu National Forest Park in Wulan County (HL, annual precipitation 217 mm), Qushigang in Dulan County (QS, 281 mm) and Hebei Forest Farm in Tongde County (HB, 470 mm). The correlation and response analysis at seasonal and extreme climate year scales were used to examine the spatial variations of the growth-climate relationship of S. przewalskii at different timberlines. Important findings Our results do not support the hypothesis that water is the main limiting factor of tree growth at the arid alpine timberline. The effect of precipitation on the radial growth of S. przewalskii were consistent across all three sampling sites, while the effects of temperature were different across sites. At HL site (low precipitation), the winter and summer minimum temperature were the main limiting factor of S. przewalskii radial growth, and this relationship did not significantly change in different extreme climate years. At QS site (middle precipitation), the radial growth of S. przewalskii was mainly limited by the minimum temperature in spring and summer, but its effect was weaker than that at low precipitation site. At HB site (high precipitation), the spring temperature had a significant negative effect on tree growth, and the positive effect of spring precipitation on tree growth was significantly enhanced in comparison with those at low and middle precipitation sites, especially in extreme high temperature and drought years. Summer precipitation did not significantly affect tree growth at high precipitation site. Our results did not support the hypothesis that the radial growth of trees at alpine timberline in arid/humid area is mainly limited by water/temperature. However, precipitation at timberline will affect the relationship between tree growth and temperature at different seasons. With the warming and humidification of the northeastern Qinghai-Xizang Plateau, the climatic limiting factors of tree growth in different timberline areas may be complicated. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

干扰对高山林线再形成过程的影响

高山林线是一类典型的生态交错带,因其特殊的结构和功能以及对外界环境的高度敏感性而成为全球气候变化研究的热点之一.本文简要介绍了高山林线的相关概念及其界定,从高山林线海拔位置波动、植被格局变化、生态交错带物种组成变化及其生理生态特征变化等几个方面阐述了干扰对高山林线再形成过程的不同影响,总结了高山林线物种对干扰的两种基本响应方式,即退行和入侵.认为人为干扰在一定程度上弱化了当前气候变暖对高山林线波动的影响,因而在不同地区必须紧密结合当地可能的干扰来讨论高山林线的波动,否则结果有可能因误差较大而失去应有的价值.指出该研究在高海拔地区进行植被恢复的指导意义.     

The interacting effects of temperature,ground disturbance,and herbivory on seedling establishment: implications for treeline advance with climate warming

Thermal control of treeline position is mediated by local environmental and ecological factors, making trends in treeline migration difficult to extrapolate geographically. We investigated the ecological dynamics of conifer establishment at treeline in the Mealy Mountains (Labrador, Canada) and the potential for its expansion with climate warming. Available seedbed and tree seedling emergence in the treeline ecotone were monitored, and seeds and seedlings of Picea mariana were planted along an elevational gradient from open-canopy forest through tree islands to alpine tundra. Experimental treatments included passive warming of daytime air, ground disturbance, and vertebrate herbivore exclosures. Responses in seed germination and seedling growth, damage, and mortality were monitored over two growing seasons, and re-surveyed after 5 years. While no tree seedlings were observed growing naturally above the treeline, planted seeds were able to germinate, develop and overwinter, and persist for 4 years in all habitats examined. Disturbance of the seedbed was important for seedling emergence in the forest and tree islands. While temperature enhancement alone had little impact on emergence, even moderate temperature increases had significantly disproportionate effects on emergence of seedlings in the alpine habitat when combined with soil disturbance, indicating that future climate warming could lead to treeline advance if viable seed and suitable substrate for recruitment are available. The positive effect of excluding herbivores suggests that herbivory may be an important filter modifying future species distribution. While seedbed conditions and herbivory would control the rate of individual species advance, the results indicate potential upslope migration of the treeline in the Mealy Mountains, with consequent loss of alpine ecosystems.     

高山林线与气候变化关系研究进展

20世纪全球气候经历了异常的变化。20世纪是过去1000年中增暖最大的1个世纪,并且90年代是最暖的10年。作为两个生态系统的过渡地带,生态过渡带是监测全球变化的重要地点,而森林和苔原之间的高山林线是全球变化最为敏感的地点。从高山林线树木个体对气候变化的响应、气候变化下林线处树木的更新、林线格局变化以及高山林线与气候变化关系研究中所采用的研究方法等方面,综合论述了国内外的研究进展,最后提出了高山林线研究中需要注意的问题,并对今后的研究趋势作了展望。     

Asymmetric effects of daytime and nighttime warming on alpine treeline recruitment

Trees at their upper range limits are highly sensitive to climate change, and thus alpine treelines worldwide have changed their recruitment patterns in response to climate warming. However, previous studies focused only on daily mean temperature, neglecting the asymmetric influences of daytime and nighttime warming on recruitments in alpine treelines. Here, based on the compiled dataset of tree recruitment series from 172 alpine treelines across the Northern Hemisphere, we quantified and compared the different effects of daytime and nighttime warming on treeline recruitment using four indices of temperature sensitivity, and assessed the responses of treeline recruitment to warming-induced drought stress. Our analyses demonstrated that even in different environmental regions, both daytime and nighttime warming could significantly promote treeline recruitment, and however, treeline recruitment was much more sensitive to nighttime warming than to daytime warming, which could be attributable to the presence of drought stress. The increasing drought stress primarily driven by daytime warming rather than by nighttime warming would likely constrain the responses of treeline recruitment to daytime warming. Our findings provided compelling evidence that nighttime warming rather than daytime warming could play a primary role in promoting the recruitment in alpine treelines, which was related to the daytime warming-induced drought stress. Thus, daytime and nighttime warming should be considered separately to improve future projections of global change impacts across alpine ecosystems.     

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。     

长白山北坡林线岳桦种群空间分布格局

许多自然林线具有的趋同特征之一即乔木树种高生长受限,常演化为矮曲状或类似于灌木的形态(即树种的灌木型)占据高山植被带,因此研究林线树种乔木型与灌木型的结构、功能差异有助于进一步理解林线形成的原因。种群分布格局作为种群相对位置定量化描述的基本特征,可以表征物种对环境适应性选择的结果,反应生态过程的综合作用。利用点格局方法,研究长白山北坡林线岳桦种群各生活史阶段、两种生活型的分布格局,结果表明,长白山北坡林线岳桦树高生长受到限制,1.5—3.0m是一个关键的树高生长阶段;相比于老树、中树,幼苗和灌木型岳桦更为均匀,对空间的异质性选择更弱;林线岳桦发育过程中,存在一个生活型分离的重要阶段。此外,相对于老树,灌木型分布更为均匀,表明低矮、多枝这种相对紧凑的生活型更适宜在过渡带生存,乔木型岳桦和灌木型岳桦可能代表着不同的生存策略。     

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.     

A re-assessment of high elevation treeline positions and their explanation

In this review I first compile data for the worldwide position of climate-driven alpine treelines. Causes for treeline formation are then discussed with a global perspective. Available evidence suggests a combination of a general thermal boundary for tree growth, with regionally variable “modulatory” forces, including the presence of certain taxa. Much of the explanatory evidence found in the literature relates to these modulatory aspects at regional scales, whereas no good explanations emerged for the more fundamental global pattern related to temperature per se, on which this review is focused. I hypothesize that the life form “tree” is limited at treeline altitudes by the potential investment, rather than production, of assimilates (growth as such, rather than photosynthesis or the carbon balance, being limited). In shoots coupled to a cold atmosphere, meristem activity is suggested to be limited for much of the time, especially at night. By reducing soil heat flux during the growing season the forest canopy negatively affects root zone temperature. The lower threshold temperature for tissue growth and development appears to be higher than 3°C and lower than 10°C, possibly in the 5.5–7.5°C range, most commonly associated with seasonal means of air temperature at treeline positions. The physiological and developmental mechanisms responsible have yet to be analyzed. Root zone temperature, though largely unknown, is likely to be most critical. Received: 3 October 1997 / Accepted: 14 April 1998     

Alpine treelines as ecological indicators of global climate change: Who has studied? What has been studied?

As the “front line” of coping with global climate change, the alpine treeline has been widely investigated by ecologists for a long time. In this study, 3180 articles in the Web of Science database from 2000 to 2021 are visualized by using CiteSpace bibliometric software from the perspectives of basic literature features (volume, discipline, journal, author, institution, and country), academic layout, research hotspot, and research frontier to have a deeper understanding of the development laws and hot spots in the alpine treeline. Results show that: (1) The number of papers published in alpine treeline research field is increasing annually, and the professional degree of papers (Sp) is 0.13–0.14, it is lower than that in other fields, implying a tendency of multi-disciplinary integration . (2) The core journals are headed by Arctic and Alpine Research, and J. Julio Camarero is the most prolific author in the field. (3) From the analysis of countries and institutions, the United States and the Chinese Academy of Sciences have leading positions in this research field. (4) Over the last 20 years, alpine treeline researchers have primarily focused on three key words (climate change, environmental characteristics, interspecific interactions, etc.), research sites (National Glacier Parks, the Alps, the Sygera Mountains, etc.), and research species (Norway spruce, European spruce, Abies georgei, etc.). (5) The first phase (2000–2009) focuses on the formation and changes of the alpine treeline in the context of climate change, and the second phase (2010−2021) focuses on the driving mechanisms of climatic factors, physiological change of tree species and grasslands on the treeline ecotones, and the influence of human activities (logging, grazing, etc.) on the alpine treeline. (6) Academic publications in this discipline have a short half-life (3.85 ± 1.70 years), and the aging rate is high and changing quickly. In the coming years, more research on alpine treeline will be devoted to explain the mechanism of tree species limitation in alpine treeline under climate change, influence of treeline changes the carbon and water cycles. There is a press need to explore the driving mechanism for treeline shift from the perspectives of plant physiology and soil carbon and nitrogen cycles.     

气候变暖对老秃顶子林线结构特征的影响

运用样带样方法和年轮气候学方法对大海林地区的气候因子和样地数据进行了分析。结果表明 ,近 30 a来老秃顶子地区气候变暖明显 ,尤其是冬季增温最明显 ,月份增温中以 2月份最大 ;寒冷时期 (12月份、翌年 1和 2月份 )和温暖时期 (6～ 9月份 )的温度都有增加 ,但是寒冷时期温度的增加幅度较大 ;冬季与夏季温差稍有减少 ,但积温有所增加 ,整年的热量正在增加。全球变暖导致的大海林地区增温对老秃顶子林线结构特征产生了很大的影响 ,由样地调查和分析可知 ,全球变暖导致林线中上部幼苗、幼树的更新和存活增多 ,森林密度加大 ,树木平均年龄降低 ,年龄结构呈倒 J字型 ,并且多呈聚集分布 ;而在林线的下部 ,幼苗更新很少 ,主要以中龄林存在 ,并且多呈零散分布形式。通过年轮分析得出 ,气候变暖导致林线树木径生长和高生长增加 ,而且增加的趋势和近 30 a来温度的变化基本一致。通过对年轮指数与气候因子的相关性分析 ,表明林线树木年轮指数与温度的相关性较强 ,而与降水的相关性较弱 ,并且年轮指数与温暖时期温度和积温呈正相关 ,而与寒冷时期温度和年平均温度呈负相关 ,表明温暖时期温度和积温控制着林线的海拔高度 ,而寒冷时期的温度和年平均温度主要对林线树种类型起着决定性的作用。从敏感度分析看出 ,林线     

高山林线形成机制及假说的探讨

高山林线作为森林向苔原过渡的敏感带,历来被生态学家视为生态脆弱区、外界干扰信号的放大器和全球变化重要的预警区.20世纪80年代开始,许多研究关注全球变化对高山林线的影响,这一阶段针对林线海拔高度、纬度位置之间的关系及对未来热环境的响应研究已成为众多学者研究的焦点之一.众多学者依据自身研究目的、不同树种出现上限的原因、局部上影响林木的因子间的差异,在区域上对高山林线的形成机制的提出了诸多假说.本文针对这些假说进行了综述和探讨,并就其中不足之处提出质疑,提出今后需要深入研究的几个方向.

Abstract：

As a sensitive transitional zone between forest and tundra, timberline has always been considered by ecologists to be an ecologically fragile zone, an amplifier of outside interference signals, and a global climate change early warning zone. Since the 1980s, many studies have been made on the effects of global climate change on alpine timberline, mainly addressed the re-lationships of the upper limits of timberline with altitude and latitude, and in particular, the re-sponses of timberline location to global warming. Several hypotheses were proposed to examine the timberline formation of various tree species affected by environmental factors. This paper sum-marized these hypotheses, and discussed some potential studies in the future.     

Climate continentality and treeline species distribution in the Alps

Abstract

The distribution of tree species and the elevation of the alpine treeline are strongly affected by climate continentality. In the present work we performed a detailed survey of the upper limits of tree vegetation in two areas with contrasting climate located in the central Italian Alps, in order to evaluate the structure of the treeline under different degrees of continentality. Tree and krummholz (stunted) individual position, their dimension and life form were recorded from the upper limit of the closed forest to the species limit. The results were compared with an estimation of tree species distribution at the treeline in the whole Lombardy Alps, performed by a survey of tree species occurrence in areas of known climatic traits. The structure of the treeline (upper limits, life form altitudinal arrangement) and its ongoing dynamics were different in the two areas: climate continentality assessed by hygric and thermal continentality indices influenced the distribution of some treeline species. Although the influence of human and geomorphologic disturbance could not be excluded, the importance of the degree of continentality must be stressed when evaluating the response of the treeline to past and present climatic change.     

Uncoupled changes in tree cover and field layer vegetation at two Pyrenean treeline ecotones over 11 years

Background: The alpine treeline ecotone is regarded as a sensor of the effects of global change on alpine plant communities. However, little is known about how treeline dynamics influence the diversity and composition of alpine plant communities. Such information is necessary to forecast how ascending montane forests may affect the composition of alpine flora.

Aims: We analysed the temporal variations in tree cover, plant diversity and composition, and the effect of tree cover dynamics on field layer vegetation over a period of 11 years, at two alpine treeline ecotones in the central Pyrenees, Spain.

Methods: Tree and field layer vegetation was sampled in permanent transects in 1998 and 2009, using the point-intercept method. Temporal changes in tree cover, plant species richness and abundance were characterised along the ecotone by using a randomisation approach, rarefaction curves, and a non-parametric multivariate test, respectively.

Results: Tree cover increased significantly at one of the sites, whereas plant species richness only increased at the other site where tree cover had not changed. Vegetation composition changed significantly at both sites, but it was not spatially coupled with changes in tree cover along the ecotone.

Conclusions: A change of tree cover does not necessarily trigger changes in the ground flora at the treeline over relatively short periods (decade scale). The results challenge our ability to infer short-term biodiversity impacts from upslope advance of forests. Integrated tree and field layer monitoring approaches are necessary to produce a better understanding of the impact of ongoing global change on treeline ecotones.     

Vegetation patterns at the alpine treeline ecotone: the influence of tree cover on abrupt change in species composition of alpine communities

Aims: The upper elevation limit of forest vegetation in mountain ranges (the alpine treeline ecotone) is expected to be highly sensitive to global change. Treeline shifts and/or ecotone afforestation could cause fragmentation and loss of alpine habitat, and are expected to trigger considerable alterations in alpine vegetation. We performed an analysis of vegetation structure at the treeline ecotone to evaluate whether distribution of the tree population determines the spatial pattern of vegetation (species composition and diversity) across the transition from subalpine forest to alpine vegetation. Location: Iberian eastern range of the Pyrenees. Methods: We studied 12 alpine Pinus uncinata treeline ecotones. Rectangular plots ranging from 940 to 1900 m² were placed along the forest‐alpine vegetation transition, from closed forest to the treeless alpine area. To determine community structure and species distribution in the treeline ecotone, species variation along the forest‐alpine vegetation transition was sampled using relevés of 0.5 m² set every 2 m along the length of each plot. Fuzzy C‐means clustering was performed to assess the transitional status of the relevés in terms of species composition. The relation of P. uncinata canopy cover to spatial pattern of vegetation was evaluated using continuous wavelet transform analysis. Results: Vegetation analyses revealed a large degree of uniformity of the subalpine forest between all treeline ecotone areas studied. In contrast, the vegetation mosaic found upslope displayed great variation between sites and was characterized by abrupt changes in plant community across the treeline ecotone. Plant richness and diversity significantly increased across the ecotone, but tree cover and diversity boundaries were not spatially coincident. Conclusions: Our results revealed that no intermediate communities, in terms of species composition, are present in the treeline ecotone. Ecotone vegetation reflected both bedrock type and fine‐scale heterogeneity at ground level, thereby reinforcing the importance of microenvironmental conditions for alpine community composition. Tree cover did not appear to be the principal driver of alpine community changes across the treeline ecotone. Microenvironmental heterogeneity, together with effects of past climatic and land‐use changes on ecotone vegetation, may weaken the expected correlation between species distribution and vegetation structure.