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This article discusses the concept of treeline dynamics in response to global warming. The authors argue that treeline ecotones in alpine and arctic settings are less sensitive to anthropogenic warming than commonly assumed. They propose the concepts of range-pinning and Allee effects as factors that may explain the reduced temperature sensitivity of treeline positions. The authors suggest that future treeline studies should consider eco-physiological and biochemical insights, empirical evidence for Allee effects, and interdisciplinary approaches to advance knowledge in biogeography and ecology. [Extracted from the article]

Mountain treelines are thought to be sensitive to climate change. However, how climate impacts mountain treelines is not yet fully understood as treelines may also be affected by other human activities. Here, we focus on "closed‐loop" mountain treelines (CLMT) that completely encircle a mountain and are less likely to have been influenced by human land‐use change. We detect a total length of ~916,425 km of CLMT across 243 mountain ranges globally and reveal a bimodal latitudinal distribution of treeline elevations with higher treeline elevations occurring at greater distances from the coast. Spatially, we find that temperature is the main climatic driver of treeline elevation in boreal and tropical regions, whereas precipitation drives CLMT position in temperate zones. Temporally, we show that 70% of CLMT have moved upward, with a mean shift rate of 1.2 m/year over the first decade of the 21st century. CLMT are shifting fastest in the tropics (mean of 3.1 m/year), but with greater variability. Our work provides a new mountain treeline database that isolates climate impacts from other anthropogenic pressures, and has important implications for biodiversity, natural resources, and ecosystem adaptation in a changing climate. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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.

Understanding how climate warming will affect the demographic rates of different ecotypes is critical to predicting shifts in species distributions. Here, we present results from a common garden, climate change experiment in which we measured seedling recruitment of lodgepole pine, a widespread North American conifer that is also planted globally. Seeds from a low-elevation provenance had more than three-fold greater recruitment to their third year than seeds from a high-elevation provenance across sites within and above its native elevation range and across climate manipulations. Heating halved recruitment to the third year of both low- and high-elevation seed sources across the elevation gradient, while watering more than doubled recruitment, alleviating some of the negative effects of heating. Demographic models based on recruitment data from the climate manipulations and long-term observations of adult populations revealed that heating could effectively halt modeled upslope range expansion except when combined with watering. Simulating fire and rapid postfire forest recovery at lower elevations accelerated lodgepole pine expansion into the alpine, but did not alter final abundance rankings among climate scenarios. Regardless of climate scenario, greater recruitment of low-elevation seeds compensated for longer dispersal distances to treeline, assuming colonization was allowed to proceed over multiple centuries. Our results show that ecotypes from lower elevations within a species' range could enhance recruitment and facilitate upslope range shifts with climate change.

Poleward shifts in species distributions are expected and frequently observed with a warming climate. In Arctic ecosystems, the strong warming trends are associated with increasing greenness and shrubification. Vertebrate herbivores have the potential to limit greening and shrub advance and expansion on the tundra, posing the question of whether changes in herbivore communities could partly mediate the impacts of climate warming on Arctic tundra. Therefore, future changes in the herbivore community in the Arctic tundra will depend on whether the community tracks the changing climates directly (i.e. occurs in response to temperature) or indirectly, in response to vegetation changes (which can be modified by trophic interactions). In this study, we used biogeographic and remotely sensed data to quantify spatial variation in vertebrate herbivore communities across the boreal forest and Arctic tundra biomes. We then tested whether present-day herbivore community structure is determined primarily by temperature or vegetation. We demonstrate that vertebrate herbivore communities are significantly more diverse in the boreal forest than in the Arctic tundra in terms of species richness, phylogenetic diversity and functional diversity. A clear shift in community structure was observed at the biome boundary, with stronger northward declines in diversity in the Arctic tundra. Interestingly, important functional traits characterizing the role of herbivores in limiting tundra vegetation change, such as body mass and woody plant feeding, did not show threshold changes across the biome boundary. Temperature was a more important determinant of herbivore community structure across these biomes than vegetation productivity or woody plant cover. Thus, our study does not support the premise that herbivore-driven limitation of Arctic tundra shrubification or greening would limit herbivore community change in the tundra. Instead, borealization of tundra herbivore communities is likely to result from the direct effect of climate warming. K E YWORDS Arctic tundra, biome boundary, boreal forest, [ABSTRACT FROM AUTHOR]

Lakes at high altitude and latitude are typically unproductive ecosystems where external factors outweigh the relative importance of in-lake processes, making them ideal sentinels of climate change. Climate change is inducing upward vegetation shifts at high altitude and latitude regions that translate into changes in the pools of soil organic matter. Upon mobilization, this allochthonous organic matter may rapidly alter the composition and function of lake bacterial communities. Here, we experimentally simulate this potential climate-change effect by exposing bacterioplankton of two lakes located above the treeline, one in the Alps and one in the subarctic region, to soil organic matter from below and above the treeline. Changes in bacterial community composition, diversity and function were followed for 72 h. In the subarctic lake, soil organic matter from below the treeline reduced bulk and taxon-specific phosphorus uptake, indicating that bacterial phosphorus limitation was alleviated compared to organic matter from above the treeline. These effects were less pronounced in the alpine lake, suggesting that soil properties (phosphorus and dissolved organic carbon availability) and water temperature further shaped the magnitude of response. The rapid bacterial succession observed in both lakes indicates that certain taxa directly benefited from soil sources. Accordingly, the substrate uptake profiles of initially rare bacteria (copiotrophs) indicated that they are one of the main actors cycling soil-derived carbon and phosphorus. Our work suggests that climate-induced changes in soil characteristics affect bacterioplankton community structure and function, and in turn, the cycling of carbon and phosphorus in high altitude and latitude aquatic ecosystems. [ABSTRACT FROM AUTHOR]

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

Mountain vegetation is strongly affected by temperature and is expected to shift upwards with climate change. Dynamic vegetation models are often used to assess the impact of climate on vegetation and model output can be compared with paleobotanical data as a reality check. Recent paleoecological studies have revealed regional variation in the upward shift of timberlines in the Northern and Central European Alps in response to rapid warming at the Younger Dryas/Preboreal transition ca. 11 700 years ago, probably caused by a climatic gradient across the Alps. This contrasts with previous studies that successfully simulated the early Holocene afforestation in the (warmer) Central Alps with a chironomid-inferred temperature reconstruction from the (colder) Northern Alps. We use LandClim, a dynamic landscape vegetation model to simulate mountain forests under different temperature, soil and precipitation scenarios around Iffigsee (2065 m a.s.l.) a lake in the Northwestern Swiss Alps, and compare the model output with the paleobotanical records. The model clearly overestimates the upward shift of timberline in a climate scenario that applies chironomid-inferred July-temperature anomalies to all months. However, forest establishment at 9800 cal. BP at Iffigsee is successfully simulated with lower moisture availability and monthly temperatures corrected for stronger seasonality during the early Holocene. The model-data comparison reveals a contraction in the realized niche of Abies alba due to the prominent role of anthropogenic disturbance after ca. 5000 cal. BP, which has important implications for species distribution models ( SDMs) that rely on equilibrium with climate and niche stability. Under future climate projections, LandClim indicates a rapid upward shift of mountain vegetation belts by ca. 500 m and treeline positions of ca. 2500 m a.s.l. by the end of this century. Resulting biodiversity losses in the alpine vegetation belt might be mitigated with low-impact pastoralism to preserve species-rich alpine meadows. [ABSTRACT FROM AUTHOR]

Circumpolar expansion of tall shrubs and trees into Arctic tundra is widely thought to be occurring as a result of recent climate warming, but little quantitative evidence exists for northern Siberia, which encompasses the world's largest forest-tundra ecotonal belt. We quantified changes in tall shrub and tree canopy cover in 11, widely distributed Siberian ecotonal landscapes by comparing very high-resolution photography from the Cold War-era 'Gambit' and 'Corona' satellite surveillance systems (1965-1969) with modern imagery. We also analyzed within-landscape patterns of vegetation change to evaluate the susceptibility of different landscape components to tall shrub and tree increase. The total cover of tall shrubs and trees increased in nine of 11 ecotones. In northwest Siberia, alder ( Alnus) shrubland cover increased 5.3-25.9% in five ecotones. In Taymyr and Yakutia, larch ( Larix) cover increased 3.0-6.7% within three ecotones, but declined 16.8% at a fourth ecotone due to thaw of ice-rich permafrost. In Chukotka, the total cover of alder and dwarf pine ( Pinus) increased 6.1% within one ecotone and was little changed at a second ecotone. Within most landscapes, shrub and tree increase was linked to specific geomorphic settings, especially those with active disturbance regimes such as permafrost patterned-ground, floodplains, and colluvial hillslopes. Mean summer temperatures increased at most ecotones since the mid-1960s, but rates of shrub and tree canopy cover expansion were not strongly correlated with temperature trends and were better correlated with mean annual precipitation. We conclude that shrub and tree cover is increasing in tundra ecotones across most of northern Siberia, but rates of increase vary widely regionally and at the landscape scale. Our results indicate that extensive changes can occur within decades in moist, shrub-dominated ecotones, as in northwest Siberia, while changes are likely to occur much more slowly in the highly continental, larch-dominated ecotones of central and eastern Siberia. [ABSTRACT FROM AUTHOR]

Russia's boreal (taiga) biome will likely contract sharply and shift northward in response to 21st century climatic change, yet few studies have examined plant response to climatic variability along the northern margin. We quantified climate dynamics, trends in plant growth, and growth-climate relationships across the tundra shrublands and Cajander larch ( Larix cajanderi Mayr.) woodlands of the Kolyma river basin (657 000 km2) in northeastern Siberia using satellite-derived normalized difference vegetation indices ( NDVI), tree ring-width measurements, and climate data. Mean summer temperatures ( T s) increased 1.0 °C from 1938 to 2009, though there was no trend ( P > 0.05) in growing year precipitation or climate moisture index ( CMI gy). Mean summer NDVI ( NDVIs) increased significantly from 1982 to 2010 across 20% of the watershed, primarily in cold, shrub-dominated areas. NDVIs positively correlated ( P < 0.05) with T s across 56% of the watershed ( r = 0.52 ± 0.09, mean ± SD), principally in cold areas, and with CMI gy across 9% of the watershed ( r = 0.45 ± 0.06), largely in warm areas. Larch ring-width measurements from nine sites revealed that year-to-year (i.e., high-frequency) variation in growth positively correlated ( P < 0.05) with June temperature ( r = 0.40) and prior summer CMI ( r = 0.40) from 1938 to 2007. An unexplained multi-decadal (i.e., low-frequency) decline in annual basal area increment ( BAI) occurred following the mid-20th century, but over the NDVI record there was no trend in mean BAI ( P > 0.05), which significantly correlated with NDVIs ( r = 0.44, P < 0.05, 1982-2007). Both satellite and tree-ring analyses indicated that plant growth was constrained by both low temperatures and limited moisture availability and, furthermore, that warming enhanced growth. Impacts of future climatic change on forests near treeline in Arctic Russia will likely be influenced by shifts in both temperature and moisture, which implies that projections of future forest distribution and productivity in this area should take into account the interactions of energy and moisture limitations. [ABSTRACT FROM AUTHOR]

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest-tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest-tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest-tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest-tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest-tundra ecotone. [ABSTRACT FROM AUTHOR]

The frequency of freezing events during the early growing season and the vulnerability to freezing of plants in European high-altitude environments could increase under future atmospheric and climate change. We tested early growing season freezing sensitivity in 10 species, from four plant functional types (PFTs) spanning three plant growth forms (PGFs), from a long-term in situ CO2 enrichment (566 vs. 370 ppm) and 2-year soil warming (+4 K) experiment at treeline in the Swiss Alps (Stillberg, Davos). By additionally tracking plant phenology, we distinguished indirect phenology-driven CO2 and warming effects from direct physiology-related effects on freezing sensitivity. The freezing damage threshold (lethal temperature 50) under ambient conditions of the 10 treeline species spanned from −6.7±0.3 °C ( Larix decidua) to −9.9±0.6 °C ( Vaccinium gaultherioides). PFT, but not PGF, explained a significant amount of this interspecific variation. Long-term exposure to elevated CO2 led to greater freezing sensitivity in multiple species but did not influence phenology, implying that physiological changes caused by CO2 enrichment were responsible for the effect. The elevated CO2 effect on freezing resistance was significant in leaves of Larix, Vaccinium myrtillus, and Gentiana punctata and marginally significant in leaves of Homogyne alpina and Avenella flexuosa. No significant CO2 effect was found in new shoots of Empetrum hermaphroditum or in leaves of Pinus uncinata, Leontodon helveticus, Melampyrum pratense, and V. gaultherioides. Soil warming led to advanced leaf expansion and reduced freezing resistance in V. myrtillus only, whereas Avenella showed greater freezing resistance when exposed to warming. No effect of soil warming was found in any of the other species. Effects of elevated CO2 and soil warming on freezing sensitivity were not consistent within PFTs or PGFs, suggesting that any future shifts in plant community composition due to increased damage from freezing events will likely occur at the individual species level. [ABSTRACT FROM AUTHOR]

The frequency of freezing events during the early growing season and the vulnerability to freezing of plants in European high-altitude environments could increase under future atmospheric and climate change. We tested early growing season freezing sensitivity in 10 species, from four plant functional types (PFTs) spanning three plant growth forms (PGFs), from a long-term in situ CO2 enrichment (566 vs. 370 ppm) and 2-year soil warming (+4 K) experiment at treeline in the Swiss Alps (Stillberg, Davos). By additionally tracking plant phenology, we distinguished indirect phenology-driven CO2 and warming effects from direct physiology-related effects on freezing sensitivity. The freezing damage threshold (lethal temperature 50) under ambient conditions of the 10 treeline species spanned from −6.7±0.3 °C ( Larix decidua) to −9.9±0.6 °C ( Vaccinium gaultherioides). PFT, but not PGF, explained a significant amount of this interspecific variation. Long-term exposure to elevated CO2 led to greater freezing sensitivity in multiple species but did not influence phenology, implying that physiological changes caused by CO2 enrichment were responsible for the effect. The elevated CO2 effect on freezing resistance was significant in leaves of Larix, Vaccinium myrtillus, and Gentiana punctata and marginally significant in leaves of Homogyne alpina and Avenella flexuosa. No significant CO2 effect was found in new shoots of Empetrum hermaphroditum or in leaves of Pinus uncinata, Leontodon helveticus, Melampyrum pratense, and V. gaultherioides. Soil warming led to advanced leaf expansion and reduced freezing resistance in V. myrtillus only, whereas Avenella showed greater freezing resistance when exposed to warming. No effect of soil warming was found in any of the other species. Effects of elevated CO2 and soil warming on freezing sensitivity were not consistent within PFTs or PGFs, suggesting that any future shifts in plant community composition due to increased damage from freezing events will likely occur at the individual species level. [ABSTRACT FROM AUTHOR]

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

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. [ABSTRACT FROM AUTHOR]

SummaryUnderstanding the response of terrestrial ecosystems to climatic warming is a challenge because of the complex interactions of climate, disturbance, and recruitment across the landscape. We use a spatially explicit model (ALFRESCO) to simulate the transient response of subarctic vegetation to climatic warming on the Seward Peninsula (80 000 km2) in north-west Alaska. Model calibration efforts showed that fire ignition was less sensitive than fire spread to regional climate (temperature and precipitation). In the model simulations a warming climate led to slightly more fires and much larger fires and expansion of forest into previously treeless tundra. Vegetation and fire regime continued to change for centuries after cessation of the simulated climate warming. Flammability increased rapidly in direct response to climate warming and more gradually in response to climate-induced vegetation change. In the simulations warming caused as much as a 228% increase in the total area burned per decade, leading to an increasingly early successional and more homogenous deciduous forest-dominated landscape. A single transient 40-y drought led to the development of a novel grassland–steppe ecosystem that persisted indefinitely and caused permanent increases in fires in both the grassland and adjacent vegetation. These simulated changes in vegetation and disturbance dynamics under a warming climate have important implications for regional carbon budgets and biotic feedbacks to regional climate. [ABSTRACT FROM AUTHOR]