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.     

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.     

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

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

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.     

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.     

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

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

高山林线形成机理研究进展

高山林线是郁闭森林与高山植被之间的分布界限,作为重要的生态过渡带,对全球和区域性气候变化的反应极为敏感,被认为是气候变化的理想监测器.高山林线研究从最初的形态描述到林线成因假说都是为了寻找高山林线形成的原因.迄今出现的高山林线成因假说都能够在局地尺度解释高山林线成因,但仍然缺乏可以普遍解释全球高山林线现象的假说.温度是林线分布的限制因子,低温限制了林线树种的生存及生长,但是低温影响了哪一个生化过程仍不明确,其影响机理还需进一步探讨.本文对高山林线形成机理,特别是对低温对高山林线植物光合特性、养分特征、非结构性碳水化合物和抗氧化系统的影响等研究进展进行综述,并提出了未来林线研究应该关注的问题.     

High-Andean vegetation and environmental gradients in northwestern Patagonia,Argentina

Abstract. We analysed the heterogeneity of high-elevation vegetation on three mountains along a west-east transect at 41 °S lat. in the Andes of northwestern Patagonia, Argentina. In this area, high-Andean vegetation occurs as islands on mountain tops above Nothofagus pumilio forests ? with the timberline at ca. 1700 m a.s.l. We recorded floristic, topographic and substrate data in 166 sites stratified according to longitude, altitude, slope and aspect. Vegetation data were classified with TWINSPAN and ordinated with Detrended Correspondence Analysis. The relationship between environmental and floristic variation was analysed using Canonical Correspondence Analysis. In order of importance, geographical longitude, altitude and aspect were the major determinants of vegetation variation, whereas substrate texture and slope appeared less important. The combination of these factors resulted in two main vegetation gradients. The first gradient is related to a moisture availability gradient, primarily determined by longitude and secondarily to variation in wind exposure (east vs. west aspect). The second vegetation gradient is related to variation in temperature, primarily determined by altitude, and secondarily by variation in insolation related to the contrast between north and south aspects. The four communities obtained with TWINSPAN are therefore associated with four characteristic habitat types: moist-cold, moist-warm, dry-cold and dry-warm. The community of warm and dry environments is the richest and has elements in common with dry steppe communities situated at lower elevations to the east, while the vegetation of the cold-moist habitat type has unique elements that are typical of the southern Andes. Although current climatic factors appear to be the major determinants of high-Andean vegetation gradients, historical events of Pleistocene times probably also affected the vegetation patterns we see today.     

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

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

Plant species distribution across two contrasting treeline ecotones in the Spanish Pyrenees

We describe the structure of two contrasting (elevation, topography,climate, vegetation, soil) alpine forest–pasture ecotones located in theCentral Pyrenees (sites Ordesa, O, and Tessó, T). We define ecotonestructure as the spatial distribution of trees of different size classes andgrowth-forms and the relationship between these aspects and the spatialdistribution of understory vegetation and substrate. The studied ecotones aredominated by Pinus uncinata Ram. and have been littleaffected by anthropogenic disturbances (logging, grazing) during this century.One rectangular plot (30 × 140 m) was located within eachsite with its longest side parallel to the slope and encompassing treeline andtimberline. The distribution of size and growth-form classes at site O followeda clear sequence of increasing size downslope from shrubby multistemmedkrummholz individuals to bigger arborescent trees. At site O, regeneration wasconcentrated near the krummholz area and over rocky substrates. This suggeststhat krummholz may modify microenvironment conditions and increase seedlingsurvival. At site T, regeneration was abundant above the treeline where thecover of the dominant understory shrub (Rhododendronferrugineum) decreased. In both ecotones the diversity of plants washigher above the treeline than in the forest and decreased going downslopecoinciding with the increase of P. uncinata cover. Thereduction of plant diversity appeared above the current timberline. At site O,the decrease was steep and spatially heterogeneous what may be due in part tothe edaphic heterogeneity. At site T the change was abrupt though smaller. Therelationships between the plant community and tree regeneration should be takeninto account in future ecological studies of treeline pattern.     

A novel framework for disentangling the scale‐dependent influences of abiotic factors on alpine treeline position

Low‐temperature growth limitation largely determines alpine treeline position globally, but treeline elevation also varies locally at a range of scales in response to multiple biotic and abiotic factors. In this study, we conceptualise how variability in treeline elevation is related to abiotic factors that act as thermal modifiers, physiological stressors, or disturbance agents. We then present a novel analytical framework for quantifying how abiotic factors influence treeline elevation at different spatial scales using New Zealand Nothofagus treelines as a case study. We delineated Nothofagus treelines in a GIS, along which we extracted data for treeline elevation and eight abiotic explanatory variables at 54 000 points. Each location was classified at each of five spatial scales based on nested river catchments, ranging from large regional to small hillslope catchments. We used hierarchical linear models to partition the variation in both treeline elevation and the eight abiotic variables by spatial scale, and then quantified the relationships between these at each spatial scale in turn. Nothofagus treeline elevation varied from 800–1740 m a.s.l. across New Zealand. Abiotic factors explained 82% of the variation in treeline elevation at the largest (regional) scale and 44–52% of variation at the four finer scales. Broad‐scale variation in Nothofagus treeline elevation was strongly associated with thermal modifiers, consistent with the idea that treelines coincide with a temperature‐driven, physiological limit. However, much of the finer‐scale variation in treeline elevation was explained by a combination of thermal, physiological stress‐related, and disturbance variables operating at different spatial scales. The conceptual model and analytical methods developed here provide a general framework for understanding treeline variation at different spatial scales.     

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.     

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.     

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.     

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。     

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.     

Diet induces phenotypic plasticity of Percichthys trucha (Valenciennes, 1833) (Perciformes,Percichthyidae) in Patagonia

The South American fish genus Percichthys, due to its great morphological variation, has included several nominal species throughout a long history. However, current genetic analyses signal the existence of only two extant species, Percichthys melanops Girard in Chile (west of the Andes) and P. trucha (Valenciennes) in Chile and Argentina (west and east of the Andes). Here the morphological variation of free embryos, larvae, and juveniles of P. trucha was analyzed using linear measurements and geometric morphometrics. Early morphological variation and compensatory growth were examined using sibling free embryos and larva. Morphological consequences of experimentally controlled food treatments were explored in juveniles. Our results showed individual variation in the size of the yolk-sac of free embryos and in the duration of the mixed feeding period. Phenotypic convergence of the upper jaw length from larva to juveniles and adults, and the causal relationship between diet and head shape was found to change as a consequence of controlled feeding. Embryonic, larval, and juvenile morphological variation and phenotypic plasticity observed in P. trucha pinpoint a possible cause for the shape variation observed in the wild. Phenotypic plasticity allows P. trucha to exploit different trophic resources and to occupy different habitats in low biodiversity lakes of Patagonia, being a major factor taking part in the past and present success of P. trucha.     

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

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

The inability of tropical cloud forest species to invade grasslands above treeline during climate change: potential explanations and consequences

The upper elevational range edges of most tropical cloud forest tree species and hence the ‘treeline’ are thought to be determined primarily by temperatures. For this reason, the treeline ecotone between cloud forests and the overlying grasslands is generally predicted to shift upslope as species migrate to higher elevations in response to global warming. Here, we propose that other factors are preventing tropical trees from shifting or expanding their ranges to include high elevation areas currently under grassland, resulting in stationary treelines despite rising mean temperatures. The inability of cloud forest species to invade the grasslands, a phenomenon which we refer to as the ‘grass ceiling’ effect, poses a major threat to tropical biodiversity as it will greatly increase risk of extinctions and biotic attrition in diverse tropical cloud forests. In this review, we discuss some of the natural factors, as well as anthropogenic influences, that may prevent cloud forest tree species from expanding their ranges to higher elevations. In the absence of human disturbances, tropical treelines have historically shifted up‐ and down‐slope with changes in temperature. Over time, increased human activity has limited forests to lower elevations (i.e. has depressed treelines), and often broken the equilibrium between species range limits and climate. Yet even in areas where anthropogenic influences are halted, cloud forests have not expanded to higher elevations. Despite the critical importance of understanding the distributional responses of tropical species to climate change, few studies have addressed the factors that influence treeline location and dynamics, severely hindering our ability to predict the fate of these diverse and important ecosystems.     

Leap frog in slow motion: Divergent responses of tree species and life stages to climatic warming in Great Basin subalpine forests

In response to climate warming, subalpine treelines are expected to move up in elevation since treelines are generally controlled by growing season temperature. Where treeline is advancing, dispersal differences and early life stage environmental tolerances are likely to affect how species expand their ranges. Species with an establishment advantage will colonize newly available habitat first, potentially excluding species that have slower establishment rates. Using a network of plots across five mountain ranges, we described patterns of upslope elevational range shift for the two dominant Great Basin subalpine species, limber pine and Great Basin bristlecone pine. We found that the Great Basin treeline for these species is expanding upslope with a mean vertical elevation shift of 19.1 m since 1950, which is lower than what we might expect based on temperature increases alone. The largest advances were on limber pine‐dominated granitic soils, on west aspects, and at lower latitudes. Bristlecone pine juveniles establishing above treeline share some environmental associations with bristlecone adults. Limber pine above‐treeline juveniles, in contrast, are prevalent across environmental conditions and share few environmental associations with limber pine adults. Strikingly, limber pine is establishing above treeline throughout the region without regard to site characteristic such as soil type, slope, aspect, or soil texture. Although limber pine is often rare at treeline where it coexists with bristlecone pine, limber pine juveniles dominate above treeline even on calcareous soils that are core bristlecone pine habitat. Limber pine is successfully “leap‐frogging” over bristlecone pine, probably because of its strong dispersal advantage and broader tolerances for establishment. This early‐stage dominance indicates the potential for the species composition of treeline to change in response to climate change. More broadly, it shows how species differences in dispersal and establishment may result in future communities with very different specific composition.