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.     

Forest-level analysis of stability under exploitation: depensation responses and catastrophe theory

Stability analysis of whole forests is proposed as a qualitative tool for the study of forest responses to partial or patchy harvests or mortality. Instead of modeling every tree or stand, aggregate tree biomass is modeled. In order to aggregate stands, spatial effects must be incorporated. It is shown that depensation growth responses (reduced growth at low biomass) are common because forests often modify harch environments to be more suitable for their growth. Depensation can result from thinning, partial mortality, patchy cutting, or clearcutting, depending on forest type and abiotic factors. Examples of these types of behaviors are given. Stability analysis of different growth regimes under exploitation are related to catastrophe theory and to optimal harvesting policies. Such qualitative analysis is shown to be applicable to data-poor regions such as the tropics where there is great concern over responses of forests to exploitation.     

Effects of disturbance and size structure on the regeneration process in a sub-boreal coniferous forest,northern Japan

The stand structure and disturbance history in a sub-boreal coniferous forest dominated byPicea jezoensis, Picea glehnii andAbies sachalinensis were investigated in four study plots set up in Taisetsuzan National Park, Japan. The effect of stand characteristics on the growth and mortality rates of understory trees was examined. Although all the stands showed inverse J-shape d.b.h. (diameter at breast height) distributions, the age structure and disturbance history differed amongst the stands. The stands with wide d.b.h. distribution (i.e. large CV and skewness) were more uneven-aged than those with narrow d.b.h. distribution (i.e. small CV and skewness). The disturbance-return interval based on the model of Hett and Loucks was 31 to 65 years. The gap ratio in the canopy was also different among the stands. These suggest that the variations in stand structure represent different occurrences of natural disturbances. Furthermore, the structural features such as size structure, canopy gap ratio and density of canopy trees also affected the growth dynamics of understory trees (≥2 m in height and <10 cm in diameter at breast height). The growth and mortality rates of understory trees changed with the canopy gap ratio and canopy tree density. The understory trees of stands with wide canopy d.b.h. distribution had higher growth and canopy recruitment rates than those of stands with narrow canopy d.b.h. distribution, contributing to the maintenance of continuous stand stratification. The understory trees of stands with narrow canopy d.b.h. distribution showed lower growth and higher mortality rates than those of stands with narrow canopy d.b.h. distribution, leading to the formation of a single-canopy structure. It is suggested that natural disturbance governs the regeneration process in the future by affecting the growth and mortality patterns of understory trees through the stand structure (size and age structure, canopy tree density, canopy gap ratio).     

An analytical model of stand dynamics as a function of tree growth, mortality and recruitment: the shade tolerance-stand structure hypothesis revisited

Light competition and interspecific differences in shade tolerance are considered key determinants of forest stand structure and dynamics. Specifically two main stand diameter distribution types as a function of shade tolerance have been proposed based on empirical observations. All-aged stands of shade tolerant species tend to have steeply descending, monotonic diameter distributions (inverse J-shaped curves). Shade intolerant species in contrast typically exhibit normal (unimodal) tree diameter distributions due to high mortality rates of smaller suppressed trees. In this study we explore the generality of this hypothesis which implies a causal relationship between light competition or shade tolerance and stand structure. For this purpose we formulate a partial differential equation system of stand dynamics as a function of individual tree growth, recruitment and mortality which allows us to explore possible individual-based mechanisms--e.g. light competition-underlying observed patterns of stand structure--e.g. unimodal or inverse J-shaped equilibrium diameter curves. We find that contrary to expectations interspecific differences in growth patterns can result alone in any of the two diameter distributions types observed in the field. In particular, slow growing species can present unimodal equilibrium curves even in the absence of light competition. Moreover, light competition and shade intolerance evaluated both at the tree growth and mortality stages did not have a significant impact on stand structure that tended to converge systematically towards an inverse J-shaped curves for most tree growth scenarios. Realistic transient stand dynamics for even aged stands of shade intolerant species (unimodal curves) were only obtained when recruitment was completely suppressed, providing further evidence on the critical role played by juvenile stages of tree development (e.g. the sampling stage) on final forest structure and composition. The results also point out the relevance of partial differential equations systems as a tool for exploring the individual-level mechanisms underpinning forest structure, particularly in relation to more complex forest simulation models that are more difficult to analyze and to interpret from a biological point of view.     

Long-term growth dynamics of natural forests in Hokkaido,northern Japan

Abstract. The long-term growth dynamics of natural forest stands on the island of Hokkaido were described on the basis of an analysis of data from 38 permanent plots spanning 15–22 yr. Stand structure was characterized by basal area, stem density and tree size variability. To detect trends in stand structure, regression models for recruitment rate (per ha per yr), mortality rate and the rate of change in stem density and tree size variability were developed by a stepwise method using initial basal area, stem density, tree size variability, species composition summarized by LNMDS ordination, altitude, annual mean temperature, annual precipitation, type of understorey vegetation, topography and slope aspect as candidates for predictor variables. The same analyses were conducted for basal area increment (net growth) and its components: survivor growth = basal area gain by growth of surviving individuals and mortality = basal area loss by death of individuals. Stem density remained generally unchanged; recruitment was relatively low even in very sparse stands. Stand basal area generally increased as survivor growth was approximately double the mortality. Recruitment rate was strongly affected by the presence of dwarf bamboo (Sasa spp.) vegetation on the forest floor which inhibited tree regeneration. Mortality rate was density-dependent; dense stands had higher mortality than sparse stands. Density change rate (recruitment rate - mortality rate) was, therefore, determined by both the type of understorey vegetation and stem density. Survivor growth was high in stands with high stem density and basal area. Mortality was dependent on basal area and altitude. Net basal area increment (net growth) was dependent only on stem density with other factors that influenced survivor growth and mortality omitted. Tree size variability decreased in stands with high tree size variability whereas it increased in stands with low size variability. Based on the obtained models for density change rate and net basal area increment, trajectories of stands were illustrated on a log-log diagram of stem density and basal area. The predicted differences in trajectories as affected by the understorey vegetation type indicated the importance of dwarf bamboo vegetation for forest dynamics on Hokkaido.     

Managing red pine stand structure to mitigate drought impacts

Reducing forest stand density through silvicultural thinning has demonstrated potential to mitigate drought impacts on growth; however, less has been studied on how changes in stand structure created by different thinning methods influence forest growth responses to drought. This research examined the growth responses to drought of natural-origin red pine in a long-term study contrasting thinning methods. Dendrochronological methods were used to examine growth responses during several drought events among stands where different thinning methods have been applied since 1950. Growth responses to drought were expressed as resistance (maintaining growth during drought), and resilience (regaining pre-drought growth). Results indicate that periodic thinning from above, which resulted in smaller diameters, has the potential to moderate drought-induced growth reductions. Larger tree diameters negatively influenced tree-level resistance and resilience across all treatments; however, the proportion of dominant trees in a stand had contrasting effects on stand-level drought responses. Stands thinned from above exhibited more complex vertical structure and increased stand-level resistance and resilience to drought-induced growth declines because competition is more stratified among smaller diameter trees. Opposite trends were observed in stands thinned from below, where the larger diameters and monolayered structure create greater competition among trees of similar size and crown position. The results of this study highlight the utility in managing for greater structural diversity to mitigate the negative effects of drought in red pine forest ecosystems.     

Assessing the potential suitability of forest stands as Kirengeshoma koreana habitat using MaxEnt

The objective of this study was to assess the potential suitability of forest stands of the Mt. Baek-un region in South Korea as habitats for Kirengeshoma koreana by determining essential biotic and abiotic environmental factors using MaxEnt. Presence data were collected from 16 plots in the study area, and a forest stand map was used to assess the potential suitability of the plot as habitat for the species. The topographic site characteristics were analyzed using GIS, and the terrain relief conditions were measured using the topographic position index. The site environmental factors that significantly influenced the potential suitability of the forest stands as habitats for K. koreana were chosen using MaxEnt. The results indicated that landform types, soil depth, and water and light availability at the forest floor were the factors that most strongly influenced K. koreana habitat suitability. These significant environmental factors were assessed to determine the forest stand sites that were most suitable as potential habitats for the species in the study area.     

Mortality of developing floodplain forests subjected to a drying climate and water extraction

River regulation and water extraction have altered the hydrology of rivers resulting in substantial changes to forest structure and the dieback of floodplain forests globally. Forest mortality, due to water extraction, is likely to be exacerbated by climate change-induced droughts. In 1965, a plantation trial was established within a natural floodplain forest to examine the effect of planting density on timber production. We used data from this trial to investigate the effect of initial stand density on the structure and dynamics of Eucalyptus camaldulensis (Dehnh.) forests. Highest density stands (8000 trees ha⁻¹) were dominated by many slender trees, mostly<10 cm in diameter, whereas the lowest density stands produced size distributions with a wider range of stem diameters and higher mean and maximum stem diameter. After 1996, the study area experienced a sharp decline in water availability due to a substantial lowering of the water table, reduced flooding frequency, a pronounced rainfall deficit and increased maximum temperatures. The drought coincided with a dramatic increase in mortality in the high-density stands, yet remained little changed in low-density treatments. Our results highlight the importance of initial stand density as a key determinant of the development of forest structure. Early thinning of high-density stands is one component of a broader management approach to mitigate impacts of human-induced drought and water extraction on developing floodplain forests.     

Red Spruce Stand Dynamics, Simulations, and Restoration Opportunities in the Central Appalachians

Red spruce (Picea rubens)–dominated forests occupied as much as 600,000 ha in West Virginia prior to exploitive logging era of the late nineteenth and early twentieth centuries. Subsequently, much of this forest type was converted to northern hardwoods. As an important habitat type for a number of rare or sensitive species, only about 12,000 ha of red spruce forests presently remain in the state. In order to assess the prospects for restoration, we examined six northern hardwood stands containing understory red spruce to (1) characterize stand dynamics and regeneration patterns and (2) simulate the effectiveness of restoration silviculture to enhance red spruce overstory recruitment. Stands originated in the late 1800s to early 1900s and are currently in the (late) stem exclusion or understory reinitiation stages. Five of the six stands had even‐aged overstories that originated after clear‐cutting. Tree‐ring chronologies show high initial growth rates consistent with stand initiation. One stand, partially harvested in 1915, was uneven aged with older, legacy residuals in the canopy. Most stands had two cohorts of understory red spruce, with more than 40% of these individuals showing prior release. Our 100‐year growth simulation suggested that a 50% basal area thinning from above could double red spruce basal area to support a mixed spruce–hardwood stand in approximately 20–40 years. These results indicate that restoration silviculture could be an effective tool for increasing the amount and quality of this reduced forest type in the central Appalachians.     

Structural characteristics and canopy dynamics of Tsuga canadensis in forests of the southern Appalachian Mountains, USA

Tsuga canadensis (L.) Carr. forests of the southern Appalachian Mountains are currently facing imminent decline induced by a nonnative insect pest, the hemlock woolly adelgid (Adelges tsugae Annand). To effectively manage these forest systems now and in the future, land managers need baseline data on forest structure and dynamics prior to large-scale Tsuga canadensis mortality. Most of our knowledge concerning the dynamics of Tsuga canadensis forests comes from more northern locations such as the Great Lakes region and New England and, therefore, may not pertain to the ecological systems found within the southern Appalachian Mountains. We examined the structure and canopy dynamics of four Tsuga canadensis forest stands within the Cataloochee watershed, in the far eastern part of Great Smoky Mountains National Park (GSMNP). We characterized the environmental settings and vertical forest layers, as well as the diameter and age-structures of each Tsuga canadensis forest stand. These environmental and structural data showed that there were indeed differences between forest stands with and without successful Tsuga canadensis regeneration. The two forest stands exhibiting successful Tsuga canadensis regeneration were located above 1,000 m in elevation on well-drained, moderately steep slopes and had the greatest canopy openness. Structural data from these two forest stands indicated a history of more continuous Tsuga canadensis regeneration. We also constructed disturbance chronologies detailing the history of canopy response to disturbance events and related these to Tsuga canadensis regeneration within each forest stand. Student t-tests adjusted for unequal variances indicated significant differences in the number of release events per tree between forest stands with and without successful Tsuga canadensis regeneration. While forest stands with successful Tsuga canadensis regeneration were more frequently disturbed by minor to major canopy disturbances, events of moderate intensity were found to be most significant in terms of regeneration. These data will be of value to land managers maintaining stands of Tsuga canadensis where treatment for hemlock woolly adelgid infestation has been successful. In areas where treatment is impractical or unsuccessful, land managers will be able to use these data to restore Tsuga canadensis forests after the wave of hemlock woolly adelgid induced mortality has passed. As of August 2008, Joshua A. Kincaid will be a member of the Environmental Studies program at Shenandoah University in Winchester, Virginia, USA     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

Hurricane Disturbance Alters Secondary Forest Recovery in Puerto Rico

Land-use history and large-scale disturbances interact to shape secondary forest structure and composition. How introduced species respond to disturbances such as hurricanes in post-agriculture forest recovery is of particular interest. To examine the effects of hurricane disturbance and previous land use on forest dynamics and composition, we revisited 37 secondary forest stands in former cattle pastures across Puerto Rico representing a range of exposure to the winds of Hurricane Georges in 1998. Stands ranged from 21 to>80 yr since agricultural abandonment and were measured 9 yr posthurricane. Stem density decreased as stands aged, while basal area and species richness tended to increase. Hurricane disturbance exerted contrasting effects on stand structure, contingent on stand age. In older stands, the basal area of large trees fell, shifting to a stand structure characteristic of younger stands, while the basal area of large trees tended to rise in younger stands with increasing hurricane disturbance. These results demonstrate that large-scale natural disturbances can alter the successional trajectory of secondary forest stands recovering from human land use, but stand age, precipitation and soil series were better predictors of changes in stand structure across all study sites. Species composition changed substantially between census intervals, but neither age nor hurricane disturbance consistently predicted species composition change. However, exposure to hurricane winds tended to decrease the abundance of the introduced tree Spathodea campanulata, particularly in smaller size classes. In all sites the abundance of the introduced tree Syzygium jambos showed a declining trend, again most strongly in smaller size classes, suggesting natural thinning through succession.     

A spatial model of forest dynamics

Effects of spatial processes on temperate deciduous forest structure and dynamics were investigated with a spatial simulator derived from a forest gap model. The multi-species neighborhood model accounted for competitive interactions and endogenous disturbance in the form of small canopy gaps. Simulated and actual spatial pattern of old-growth stands were compared. The 400 yr simulations produced a pattern scale (0.07–0.2 ha patches) similar to that of an actual stand; simulated pattern intensity was greater than actual intensity, however. Distances to nearest neighbor were somewhat similar for trees in the simulated and actual stands; yet the frequency distributions of distance to nearest neighbor values differed substantially. The simulated stand patterns were generally less random than the actual patterns. Spatial pattern changed markedly during the course of simulated succession. Pattern approached a random dispersion in early succession. Intensity peaked at mid-succession (ca. 150 yr) with a hyperdispersed overstory and a strongly clumped understory. Pattern intensity diminished in late succession as a mixed size structure developed. Old-growth patch size was greater than the neighborhood (or gap) size, suggesting the gap-sized areas do not behave independently.     

Dynamics of postfire regeneration of Pinus brutia Ten. in an artificial forest ecosystem of northern Greece

This paper deals with the dynamics of postfire regeneration of Pinus brutia Ten. in an artificial forest ecosystem of North Greece, after a fire in 1982. The following issues are studied: the natural development of P. brutia stands 20 years after the fire, the current stand structure, and the influence of thinning treatment on stand population dynamics and tree growth patterns. The present work summarises and updates data taken during the years 1987–2002. The results show that the postfire regeneration was successful and contributed to the re-establishment of the pre-fire forest not only at the sites of good quality but at the medium quality sites as well. Regarding the postfire development, it is observed that an abundant P. brutia re-establishment is followed by a natural and gradual reduction of tree population caused by the influence of the physical environment during the early postfire years and caused by self-thinning later. The stands have entered the stem exclusion stage and they are growing at a narrow spacing in all cases. The evolution pattern and the stand structure were affected by thinning, which resulted in the improvement of tree quality and growth and accelerated their early fruition, thus contributing to higher ecosystem resilience.     

Tree mortality of five major species on Hokkaido Island, northern Japan

The mortality rates of five major tree species (Abies sachalinensis, Acer mono, Magnolia obovata, Quercus crispula, Tilia japonica) on Hokkaido Island, northern Japan were modeled using logistic regressions based on the long-term observation of 8929 individuals in 65 permanent plots. Individual size, recent growth, one-sided and two-sided interactions with neighbors, climatic and topographic factors were used as independent variables in the logistic regressions, and relevant variables were selected using the stepwise method. For all species analyzed, the final models significantly explained tree mortality rates. The regression analyses indicated that individual size and/or recent growth had effects on tree mortality. A significant effect of one-sided interaction on tree mortality was detected for three species, and a significant effect of two-sided interaction was detected for two species. The interactions with neighbors were not necessarily competitive. For all species analyzed, climatic and topographic factors affected tree mortality. The mortality models obtained can be used for forest dynamics simulations. One ecological question was examined using these models. Species that can grow fast in forest stands with smaller basal areas tended to have high mortality rates in forest stands with larger basal areas. Some ecological characteristics of the analyzed species are described based on the results of the regressions.     

Effects of Disturbance on Fine Root Dynamics in the Boreal Forests of Northern Ontario, Canada

Wildfire and clearcutting are two main disturbances in North American forests, but whether root systems may respond differently to such disturbances is unknown. Here, we studied how the dynamics of fine roots (<2 mm in diameter) varied with stand origins in a boreal forest in northern Ontario, Canada. Fine root biomass increased with stand age, but did not differ between stands originating from fire and clearcutting. By contrast, fine root production, mortality and turnover rates were lower in 3- and 11-year-old clearcut-origin than fire-origin stands, but did not differ in 29-year-old stands of different stand origins. The lower rates of production, mortality, and turnover rates in 3- and 11-year-old clearcut-origin than fire-origin stands are attributable to a lower density of shrubs and herbs and larger nutrient pools after clearcutting than fire. The similarities among 29-year-old stands indicate that the effects of stand origin on fine root processes tend to converge at this time scale. Our results illustrate that time scale is critical for assessing ecosystem responses to disturbances.     

SELF‐THINNING AND SIZE INEQUALITY DYNAMICS IN A CLONAL SEAWEED (SARGASSUM LAPAZEANUM,PHAEOPHYCEAE)1

Fronds of clonal seaweeds with extensive holdfasts relative to frond size are known not to self‐thin during growth, even in crowded stands. We tested whether frond self‐thinning would occur for such a seaweed since these traits are more similar to those of unitary seaweeds, which do self‐thin in crowded conditions. We used Sargassum lapazeanum Setch. et N. L. Gardner (Fucales, Phaeophyceae) from the Pacific coast of Mexico, for which we first confirmed its clonal nature by performing a regeneration experiment in culture tanks. During the growth season (winter to late spring), S. lapazeanum stand biomass increased, while frond density and size inequality (Gini coefficient for frond biomass) decreased. These results indicate that self‐thinning occurred at the frond level. We propose a conceptual model for frond dynamics for clonal seaweeds in general. In stands of clonal species with small fronds and relatively extensive holdfasts (particularly when holdfasts are perennial), frond dynamics would be determined mostly by intraclonal regulation, which seems to prevent excessive crowding from occurring. Such species display a positive biomass–density relationship during the growth season. On the contrary, in stands of clonal species with large fronds relative to holdfast size, frond dynamics would be determined mostly by interactions among genets. For such species, self‐thinning may be detected at the frond level in crowded stands, resulting in a negative biomass–density relationship during growth.     

Capturing juvenile tree dynamics from count data using Approximate Bayesian Computation

The juvenile life stage is a crucial determinant of forest dynamics and a first indicator of changes to species' ranges under climate change. However, paucity of detailed re-measurement data of seedlings, saplings and small trees means that their demography is not well understood at large scales, and rarely represented in forest models in detail. In this study we quantify the effects of climate and density dependence on recruitment and juvenile growth and mortality rates of thirteen species measured in the Spanish Forest Inventory. Single-census sapling count data is used to constrain demographic parameters of a simple forest juvenile dynamics model based on the perfect plasticity approximation model (PPA) within a likelihood-free parameterisation method, Approximate Bayesian Computation. Our results highlight marked differences between species, and the important role of climate and stand structure, in controlling juvenile dynamics. Recruitment had a hump-shaped relationship with conspecific density, and for most species conspecific competition had a stronger negative effect than heterospecific competition. Mediterranean species showed on average higher mortality and lower growth rates than temperate species, and in low density stands recruitment and mortality rates were positively correlated. Under climate change our model predicted declines in recruitment rates for almost all species. Reliable predictive models of forest dynamics should include realistic representation of critical early life-stage processes and our approach demonstrates that existing coarse count data can be used to parameterise such models. Approximate Bayesian Computation may have wide application in many fields of ecology to unlock information about past processes from single survey observations.     

Large old trees increase growth under shifting climatic constraints: Aligning tree longevity and individual growth dynamics in primary mountain spruce forests

In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with potentially dire consequences for biodiversity and carbon storage. However, the influence of tree-level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies [L.] H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed-linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks.