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。     

Multi-scale Analysis of Environmental Conditions and Conifer Seedling Distribution Across the Treeline Ecotone of Northern Manitoba, Canada

Treeline represents not only an important physiognomic boundary but also an important transition between disjunct mesoclimates and environmental limitations on establishment of tree species. The circumboreal treeline is controlled by some still to be understood physiological mechanism dependent on air temperatures, though younger life stages are increasingly influenced by numerous other biotic and abiotic factors at finer spatial and temporal scales. The goal of this study was to evaluate environmental and reproductive characteristics across treeline around Churchill, Manitoba, and to determine which factors are most important for successful seedling establishment by tamarack, white spruce and black spruce. We examined mid-winter snowpack, soil characteristics, seed viability, seedling establishment, and dominant vegetation at sites within forest and at treeline. Growing season was longer at treeline due to less snow accumulation, though soil temperatures were more variable throughout the year when compared with forest. Conifer seed germination was greater than 88% for most of the region and total seedling density was relatively consistent between sites. Seedlings were negatively associated with other plants within the forest, but low stature vegetation seemed to facilitate establishment at treeline. The associations between seedling establishment and habitat availability observed at several sites suggest that treeline advance in the Churchill area could be contingent on the facilitative effects of plants at and beyond treeline. The results of this study support the premise that fine-scale biotic and abiotic patterns and processes such as snowpack and facilitation by neighboring vegetation certainly cannot be overlooked in analyses of patterns at treeline in a changing Subarctic.     

Sensitivity and response of northern hemisphere altitudinal and polar treelines to environmental change at landscape and local scales

The sensitivity and response of northern hemisphere altitudinal and polar treelines to environmental change are increasingly discussed in terms of climate change, often forgetting that climate is only one aspect of environmental variation. As treeline heterogeneity increases from global to regional and smaller scales, assessment of treeline sensitivity at the landscape and local scales requires a more complex approach than at the global scale. The time scale (short‐, medium‐, long‐term) also plays an important role when considering treeline sensitivity. The sensitivity of the treeline to a changing environment varies among different types of treeline. Treelines controlled mainly by orographic influences are not very susceptible to the effects of warming climates. Greatest sensitivity can be expected in anthropogenic treelines after the cessation of human activity. However, tree invasion into former forested areas above the anthropogenic forest limit is controlled by site conditions, and in particular, by microclimates and soils. Apart from changes in tree physiognomy, the spontaneous advance of young growth of forest‐forming tree species into present treeless areas within the treeline ecotone and beyond the tree limit is considered to be the best indicator of treeline sensitivity to environmental change. The sensitivity of climatic treelines to climate warming varies both in the local and regional topographical conditions. Furthermore, treeline history and its after‐effects also play an important role. The sensitivity of treelines to changes in given factors (e.g. winter snow pack, soil moisture, temperature, evaporation, etc.) may vary among areas with differing climatic characteristics. In general, forest will not advance in a closed front but will follow sites that became more favourable to tree establishment under the changed climatic conditions.     

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.     

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.     

Constraints on treeline advance in a warming climate: a test of the reproduction limitation hypothesis

Aims Several mechanistic hypotheses have been developed to explain the existence of alpine treelines worldwide. The reproduction limitation hypothesis (RLH) postulates that reproductive processes such as seed production and dispersal are restricted in treeline environments, thereby limiting the establishment of new individuals in advance of extant treelines. Despite its popularity, no study has tested this hypothesis in a comprehensive fashion. In this experiment, we attempted to answer whether there are enough viable seeds being dispersed beyond treeline for sexually mediated treeline advance, and what the implications of climate change might be on these processes.Methods We established 30 plots across two aspects (north vs. south) and three elevational habitats (forest, treeline and tundra) in a white spruce (Picea glauca) boreal forest-alpine tundra ecotone in southwest Yukon, Canada. In each plot, tree characteristics, seed production and predispersal damage were measured. Additionally, eight dispersal trays were positioned in each plot to measure seed rain, and germination trials with and without predation exclosures were constructed in a subset of plots to quantify dispersal and germination success.Important findings Results were highly variable both temporally and spatially. In 2014, a mast year, 69% of adult trees produced cones compared to 0.4% in the following year. Higher density of trees in forest plots compared to treeline and tundra resulted in greater seed production at lower elevations. Across all plots, 88% of seeds were damaged before dispersal or were not viable. Treeline plots had significantly greater predispersal damage. Seed rain was greater in south-facing plots than north-facing plots. Less than 2% of seeds produced on the landscape were dispersed into Tundra plots, located 50 m above treeline. There was a net movement of seeds from the north-facing slope to the south at our study site, likely due to prevailing winds during the dispersal period. Germination counts were more than double on north-facing slopes and one-third higher inside exclosures. Cumulatively, the results provide some evidence for the RLH. Collectively, the high amount of predispersal damage and non-viable seeds, variability associated with dispersal and significant seed predation can functionally influence treeline dynamics. These findings suggest that global treeline distribution models, which rely largely on temperature, may not be entirely accurate for predicting treeline advance—at least at finer temporal scales. Many stochastic factors need to align temporally for successful advance, which is likely to result in a lag of many decades between the period of temperature amelioration and an increased number of trees beyond extant treelines.     

The importance of biotic filtering on boreal conifer recruitment at alpine treeline

Treeline, the ecotone where forest transitions to alpine or tundra ecosystems, is considered the thermal limit to tree growth and survival. Despite temperature increases across mountainous areas and high latitudes globally, there has been no ubiquitous change in treeline position. The process of range expansion must initially depend on increased recruitment at, or beyond current range limits and recruitment limitations have been hypothesized as a mechanism for the variable response of treeline position to climate warming. We conducted a unique series of observational and experimental studies to quantify early-life stage constraints, from seed production to seedling establishment, on black spruce Picea mariana and tamarack Larix laricina recruitment at a model alpine treeline in Newfoundland, Canada. We found recruitment at treeline to be simultaneously seed and establishment limited. The treeline population produced fewer seeds than the forest population and black spruce seeds produced at treeline were less viable. Tamarack was more seed limited than black spruce where seed viability was low regardless of altitudinal position. Post-dispersal seed predation greatly constrained recruitment across the altitudinal gradient; however, black spruce seeds experienced the lowest levels of invertebrate seed predation on the lichen mat at treeline. If seeds were not consumed, individuals at treeline were establishment limited where germination and seedling establishment was both less abundant and delayed on lichen substrate. Our study highlights the need for multiple factors to align temporally for significant recruitment at treeline to occur.     

A world-wide study of high altitude treeline temperatures

Aim At a coarse scale, the treelines of the world's mountains seem to follow a common isotherm, but the evidence for this has been indirect so far. Here we aim at underpinning this with facts. Location We present the results of a data‐logging campaign at 46 treeline sites between 68° N and 42° S. Methods We measured root‐zone temperatures with an hourly resolution over 1–3 years per site between 1996 and 2003. Results Disregarding taxon‐, landuse‐ or fire‐driven tree limits, high altitude climatic treelines are associated with a seasonal mean ground temperature of 6.7 °C (±0.8 SD; 2.2 K amplitude of means for different climatic zones), a surprisingly narrow range. Temperatures are higher (7–8 °C) in the temperate and Mediterranean zone treelines, and are lower in equatorial treelines (5–6 °C) and in the subarctic and boreal zone (6–7 °C). While air temperatures are higher than soil temperatures in warm periods, and are lower than soil temperatures in cold periods, daily means of air and soil temperature are almost the same at 6–7 °C, a physics driven coincidence with the global mean temperature at treeline. The length of the growing season, thermal extremes or thermal sums have no predictive value for treeline altitude on a global scale. Some Mediterranean (Fagus spp.) and temperate South Hemisphere treelines (Nothofagus spp.) and the native treeline in Hawaii (Metrosideros) are located at substantially higher isotherms and represent genus‐specific boundaries rather than boundaries of the life‐form tree. In seasonal climates, ground temperatures in winter (absolute minima) reflect local snow pack and seem uncritical. Main conclusions The data support the hypothesis of a common thermal threshold for forest growth at high elevation, but also reflect a moderate region and substantial taxonomic influence.     

Are treelines advancing? A global meta-analysis of treeline response to climate warming

Treelines are temperature sensitive transition zones that are expected to respond to climate warming by advancing beyond their current position. Response to climate warming over the last century, however, has been mixed, with some treelines showing evidence of recruitment at higher altitudes and/or latitudes (advance) whereas others reveal no marked change in the upper limit of tree establishment. To explore this variation, we analysed a global dataset of 166 sites for which treeline dynamics had been recorded since 1900 AD. Advance was recorded at 52% of sites with only 1% reporting treeline recession. Treelines that experienced strong winter warming were more likely to have advanced, and treelines with a diffuse form were more likely to have advanced than those with an abrupt or krummholz form. Diffuse treelines may be more responsive to warming because they are more strongly growth limited, whereas other treeline forms may be subject to additional constraints.     

Microsite and elevation zone effects on seed pilferage,germination, and seedling survival during early whitebark pine recruitment

Tree recruitment is a spatially structured process that may undergo change over time because of variation in postdispersal processes. We examined seed pilferage, seed germination, and seedling survival in whitebark pine to determine whether 1) microsite type alters the initial spatial pattern of seed caches, 2) higher abiotic stress (i.e. higher elevations) exacerbates spatial distribution changes, and 3) these postdispersal processes are spatially clustered. At two study areas, we created a seed distribution pattern by burying seed caches in microsite types frequently used by whitebark pine's avian seed disperser (Clark's nutcracker) in upper subalpine forest and at treeline, the latter characterized by high abiotic environmental stress. We monitored caches for two years for pilferage, germination, and seedling survival. Odds of pilferage (both study areas), germination (northern study area), and survival (southern study area) were higher at treeline relative to subalpine forest. At the southern study area, we found higher odds of 1) pilferage near rocks and trees relative to no object in subalpine forest, 2) germination near rocks relative to trees within both elevation zones, and 3) seedling survival near rocks and trees relative to no object at treeline. No microsite effects were detected at the northern study area. Findings indicated that the microsite distribution of seed caches changes with seed/seedling stage. Higher odds of seedling survival near rocks and trees were observed at treeline, suggesting abiotic stress may limit safe site availability, thereby shifting the spatial distribution toward protective microsites. Higher odds of pilferage at treeline, however, suggest rodents may limit treeline recruitment. Further, odds of pilferage were higher near rocks and trees relative to no object in subalpine forest but did not differ among microsites at treeline, suggesting pilferage can modulate the spatial structure of regeneration, a finding supported by limited clustering of postdispersal processes.     

Microtopographic Control of Treeline Advance in Noatak National Preserve, Northwest Alaska

Arctic treeline positions are of fundamental importance to the function of high latitude landscapes, as regulators of surface energy exchange and carbon cycling. Most studies aimed at explaining current and predicting future treeline positions have examined growth trends in mature treeline trees, but treeline advance requires seed production, germination, seedling establishment, and recruitment of new trees beyond the treeline and these processes may not be well correlated with growth of mature trees. Tussock tundra is a widespread, microtopographically complex vegetation type, covering vast areas of northern Alaska and Siberia. We examined a site where the white spruce (Picea glauca) treeline has recently advanced into tussock tundra and asked if white spruce seedlings (~36-cm tall) occurred in tussocks, inter-tussocks, and frost boils disproportionate to the abundance of these habitats in the treeline ecotone. We found that seedlings disproportionately occurred in tussocks and frost boils. Seedlings found in tussocks and frost boils also showed greater branch growth than those in inter-tussocks. Tussocks and frost boils had higher soil temperatures than inter-tussocks. Tussocks had higher N, P, and K availability, whereas frost boils had greater secondary and micronutrient availability. The disproportionate occurrence of seedlings in tussocks and frost boils can likely be explained by the combination of warmer soils and greater nutrient availability. It is also possible that low competition for primary nutrients contributes to the success of seedlings in frost boils. Results of the study highlight the importance of biotic and abiotic facilitation as mediators of treeline advance in a changing Arctic.     

Recent treeline dynamics are similar between dry and mesic areas of Nepal,central Himalaya

Aims We investigated the treeline dynamics of two environmentally contrasting areas in the Nepalese Himalaya to address the following questions: (i) Does the timing of establishment of the current treeline differ between the two study areas, and can area-specific treeline developments be identified? (ii) Do recruitment patterns and height growth indicate recent climate-driven treeline advance, following the general prediction for the central Himalayan region, in the two study areas?Methods A dry-climate treeline dominated by Pinus wallichiana and a mesic-climate treeline with Abies spectabilis were selected for study. In each area, we sampled the size and age structure of the study species along three elevational transects (20-m wide) from the forest line to the tree species line crossing the treeline. We also sampled treeline trees from within and outside transects to reconstruct past treeline establishment dynamics.Important findings Despite differences in moisture regimes, tree species and recent climate trends, our two study areas showed very similar treeline dynamics over the past six decades. In both areas, the recruitment of treeline trees indicates stationary treelines over the past six decades with the current treelines being dominated by trees that were established around 1990. The mesic area has experienced an overall climatic warming trend, and the stationary Abies treeline is hypothesized to be regulated by non-climatic factors, notably grazing. The dry area has not experienced warming but increased climatic variability and some very cool summers in the recent decades may explain the stationary to weakly receding Pinus treeline, which appears more climatically controlled with decreased recruitment over the past decades and decreased growth towards higher elevations. In both areas, there is a potential for treeline advance, depending on future land use and climate change. Our results highlight the importance of conducting treeline ecotone analyses for several sites or areas, and considering both climatic and non-climatic drivers of the treeline dynamics within each of these areas, for understanding regional treeline dynamics.     

High and dry: post‐fire tree seedling establishment in subalpine forests decreases with post‐fire drought and large stand‐replacing burn patches

Aim Climate warming and increased wildfire activity are hypothesized to catalyse biogeographical shifts, reducing the resilience of fire‐prone forests world‐wide. Two key mechanisms underpinning hypotheses are: (1) reduced seed availability in large stand‐replacing burn patches, and (2) reduced seedling establishment/survival after post‐fire drought. We tested for regional evidence consistent with these mechanisms in an extensive fire‐prone forest biome by assessing post‐fire tree seedling establishment, a key indicator of forest resilience. Location Subalpine forests, US Rocky Mountains. Methods We analysed post‐fire tree seedling establishment from 184 field plots where stand‐replacing forest fires were followed by varying post‐fire climate conditions. Generalized linear mixed models tested how establishment rates varied with post‐fire drought severity and distance to seed source (among other relevant factors) for tree species with contrasting post‐fire regeneration adaptations. Results Total post‐fire tree seedling establishment (all species combined) declined sharply with greater post‐fire drought severity and with greater distance to seed sources (i.e. the interior of burn patches). Effects varied among key species groups. For conifers that dominate present‐day subalpine forests (Picea engelmannii, Abies lasiocarpa), post‐fire seedling establishment declined sharply with both factors. One exception was serotinous Pinus contorta, which did not vary with either factor. For montane species expected to move upslope under future climate change (Larix occidentalis, Pseudotsuga menziesii, Populus tremuloides) and upper treeline species (Pinus albicaulis), establishment was unrelated to either factor. Greater post‐fire tree seedling establishment on cooler/wetter aspects suggested local topographic refugia during post‐fire droughts. Main conclusions If future drought and wildfire patterns manifest as expected, post‐fire tree seedling establishment of species that currently characterize subalpine forests could be substantially reduced. Compensatory increases from lower montane and upper treeline species may partially offset these reductions, but our data suggest important near‐ to mid‐term shifts in the composition and structure of high‐elevation forests under continued climate warming and increased wildfire activity.     

Effects of Climate, Site Conditions, and Seed Quality on Recent Treeline Dynamics in NW Russia: Permafrost and Lack of Reproductive Success Hamper Treeline Advance?

Treeline advance alters albedo and carbon storage and is an important feedback mechanism to the global climate system. Establishment of trees north of the treeline requires favorable climate, suitable microsites, and viable seeds. Here we studied the influence of climate and microsite conditions on tree and seedling growth at four transects from forest through woodland to tundra in NW Russia, and tested the viability of seeds from the region. General growth patterns and establishment periods of the treeline species Picea obovata are similar across the study sites suggesting a regional driver (for example, climate). Individuals established as early as the 1640s, but mainly between 1850 and 1880, and during a major and continental scale establishment wave in the 1950s and 1960s. No establishment occurred after 1982. Older trees mainly showed significant and stable positive relationships to growing year summer temperatures and significant stable negative correlations to previous year summer temperatures in nearly all plots. Trees from the last establishment wave showed more mixed responses, but current year summer temperature positively affected growth. Active layer depth was similar in all plots with trees but decreased sharply in treeless tundra. A major role for the lack of recent establishment seems to be very low seed viability, possibly combined with early strong fall frosts, which might have severely limited successful recruitment in the last decades of the twentieth century. For a successful establishment of P. obovata in tundra areas of NW Russia, permafrost degradation and (generally) warmer winters might be a prerequisite.     

High solar radiation hinders tree regeneration above the alpine treeline in northern Ecuador

Many tropical alpine treelines lie below their climatic potential, because of natural or anthropogenic causes. Forest extension above the treeline depends on the ability of trees to establish in the alpine environment. This ability may be limited by different factors, such as low temperatures, excess solar radiation, competition, soil properties, dispersal ability, and fires. In this paper we address the following two questions: Do trees regenerate above the present treeline, and what are the inhibiting factors for tree establishment? To answer these questions we described the spatial pattern of recent tree establishment below and above the present treeline in northern Ecuador. Also, we experimentally transplanted seedlings into the alpine vegetation (páramo) and the forest, and investigated the effect of shade, neighboring plants, and substrate on their survival. The number of naturally occurring tree sprouts (seedlings, saplings and ramets) was highest just outside the forest, and decreased with distance to the forest edge. However, only two species that were radiation-tolerant made up these high numbers, while other species were rare or absent in the páramo. In the forest, the species diversity of sprouts was high and the abundance per species was relatively low. The transplanted seedlings survived least in experimental plots without artificial shade where neighboring plants were removed. Seedling survival was highest in artificially shaded plots and in the forest. This shade-dependence of most tree species can strongly slow down forest expansion toward the potential climatic treeline. Due to the presence of radiation-tolerant species, the complete lack of forest expansion probably needs to be ascribed to fire. However, our results show that natural processes can also explain both the low position and the abruptness of tropical treelines.     

Competitor or facilitator? The ambiguous role of alpine grassland for the early establishment of tree seedlings at treeline

Alpine treelines are expected to move upslope with a warming climate. However, so far treelines have responded inconsistently and future shifts remain difficult to predict since many factors unrelated to temperature, such as biotic interactions, affect responses at the local scale. Especially during the earliest regeneration stages, trees can be strongly influenced by alpine vegetation via both competition and facilitation. We aimed to understand the relative importance of these two types of interaction in different vegetation structures for treeline regeneration dynamics. Effects of herbaceous alpine vegetation on seedling emergence and first‐year performance were studied in a field experiment in the French Alps (2100 m a.s.l.) with five important European treeline tree species: Larix decidua, Picea abies, Pinus cembra, Pinus uncinata and Sorbus aucuparia. Total emergence and locally‐germinated seedling survival were not affected, but for seedlings planted at two months of age, negative vegetation impacts dominated for all response parameters: first‐year survival, growth and carbohydrate accumulation. However, in the winter half‐year, evergreen tree seedlings increased carbohydrate reserves under the protection of senescent herbs. Also, responses of locally‐germinated seedlings suggest facilitative vegetation effects in the first two months after emergence. Thus, the interaction switched between competition and facilitation according to ontogenetic stage and seasons. Still, the net outcome after one year was negative, but species differed in their susceptibilities. Because initial establishment is the first bottleneck determining whether treelines remain stable or move upslope, understanding establishment, including site‐, life‐stage and species‐specific processes, is essential for understanding observed treeline spatial patterns and dynamics. When developing predictive models of treeline dynamics, all these ‘local’ aspects should be incorporated in addition to more global drivers like changes in temperature.     

Promoting survival prospects of rare plants

Many hypotheses have been put forward to explain the structure and position of alpine treelines. The spatial complexity of the ecotone, ranging from sharp boundaries to networks of tree patches within a heath matrix, may explain why no consensus has been reached. In this paper, we discuss factors from abiotic disturbances to herbivory that may help understand the spatial structure of the alpine treeline ecotone in Fennoscandia. The ecotone is dominated by mountain birch (Betula pubescens ssp. tortuosa), and may show a wide range of spatial structures. We discuss the influence of topography, seed limitations, seedling establishment, growth limitations, abiotic disturbances and herbivory as structuring factors. All of these factors may operate, but their relative importance in space and time is unknown. There is a basic difference between factors that prevent the establishment of trees, and thus act on early life history stages, and factors that thin out a previously dense forest, and thus act on adult trees. Mortality caused directly or indirectly by geometrid moths may belong to the latter category. We suggest that seedling and sapling mortality is more important than seed limitation for the establishment of new individuals in the treeline ecotone. Important mortality factors may be abiotic disturbances, competition (or allelopathy) from field layer plants and herbivory. The relative role of these factors needs to be examined further.     

Strong topographic sheltering effects lead to spatially complex treeline advance and increased forest density in a subtropical mountain region

Altitudinal treelines are typically temperature limited such that increasing temperatures linked to global climate change are causing upslope shifts of treelines worldwide. While such elevational increases are readily predicted based on shifting isotherms, at the regional level the realized response is often much more complex, with topography and local environmental conditions playing an important modifying role. Here, we used repeated aerial photographs in combination with forest inventory data to investigate changes in treeline position in the Central Mountain Range of Taiwan over the last 60 years. A highly spatially variable upslope advance of treeline was identified in which topography is a major driver of both treeline form and advance. The changes in treeline position that we observed occurred alongside substantial increases in forest density, and lead to a large increase in overall forest area. These changes will have a significant impact on carbon stocking in the high altitude zone, while the concomitant decrease in alpine grassland area is likely to have negative implications for alpine species. The complex and spatially variable changes that we report highlight the necessity for considering local factors such as topography when attempting to predict species distributional responses to warming climate.     

Seedling dynamics at elevation limits: Complex interactions beyond seed and microsite limitations

? Premise of the study: Limited availability of seeds and microsites are important constraints for seedling emergence and survival. However, little is known about population-level feedbacks between seed and microsite limitation and how these relationships vary in contrasting conditions. Focusing on Armeria caespitosa, a high-mountain endemic, we asked whether seedling establishment was simultaneously limited by seed and microsite availability, whether the balance between seed and microsite limitations varied with contrasting environments, and whether seed and microsite limitations interacted with each other. ? Methods: We studied seedling emergence and survival at the edges of the elevation range of A. caespitosa in central Spain over 4 years. We used mixed linear models and structural equation modeling. ? Key results: Our results showed that A. caespitosa recruits similar numbers of new seedlings per plot in contrasting environments. Seedling emergence and survival in A. caespitosa were both limited by seed and microsite availability. Habitat suitability (the surrogate of microsite limitation) varied between populations and indirectly affected fertility (the surrogate of seed limitation). ? Conclusions: Seedling emergence and survival are both seed and microsite limited, there are interactions between seed and microsite limitations, and both factors, which control seedling establishment, and their interconnections, vary among local populations.     

Treeline form – a potential key to understanding treeline dynamics

Aim Treelines occur globally within a narrow range of mean growing season temperatures, suggesting that low‐temperature growth limitation determines the position of the treeline. However, treelines also exhibit features that indicate that other mechanisms, such as biomass loss not resulting in mortality (dieback) and mortality, determine treeline position and dynamics. Debate regarding the mechanisms controlling treeline position and dynamics may be resolved by identifying the mechanisms controlling prominent treeline spatial patterns (or ‘form’) such as the spatial structure of the transition from closed forest to the tree limit. Recent treeline studies world‐wide have confirmed a close link between form and dynamics. Location The concepts presented refer to alpine treelines globally. Methods In this review, we describe how varying dominance of three general ‘first‐level’ mechanisms (tree performance: growth limitation, seedling mortality and dieback) result in different treeline forms, what ‘second‐level’ mechanisms (stresses, e.g. freezing damage, photoinhibition) may underlie these general mechanisms, and how they are modulated by interactions with neighbours (‘third‐level’ mechanisms). This hierarchy of mechanisms should facilitate discussions about treeline formation and dynamics. Results We distinguish four primary treeline forms: diffuse, abrupt, island and krummholz. Growth limitation is dominant only at the diffuse treeline, which is the form that has most frequently responded as expected to growing‐season warming, whereas the other forms are controlled by dieback and seedling mortality and are relatively unresponsive. Main conclusions Treeline form provides a means for explaining the current variability in treeline position and dynamics and for exploring the general mechanisms controlling the responses of treelines to climatic change. Form indicates the relative dependence of tree performance on various aspects of the external climate (especially summer warmth versus winter stressors) and on internal feedbacks, thus allowing inferences on the type as well as strength of climate‐change responses.