The potential for using Larix decidua ring widths in reconstructions of larch budmoth (Zeiraphera diniana) outbreak history: dendrochronological estimates compared with insect surveys

In the Alps, larch (Larix decidua Mill.) forests show periodic discolouration due to larch budmoth (LBM) outbreaks (Zeiraphera diniana Guénée, Lepidoptera: Tortricidae). Tree defoliation causes severe reductions in radial growth, visible in tree rings. This paper aims at reconstructing LBM outbreak history, and critically examining the potential for using dendrochronological data by comparing tree-ring estimates with insect surveys. The occurrence of LBM outbreaks was investigated using 249 cores from larch growing near the timberline in three regions of the French Alps (Briançonnais, Maurienne, and Tarentaise). Years with an abrupt decrease in radial growth (-40%) were considered as negative pointer years reflecting the potential impact of LBM. The comparison with three non-host conifers (Norway spruce, stone and mountain pines) permitted us to distinguish growth reductions in larch due to climatic effects from those due to defoliation by LBM. The dendrochronological data were matched with information reporting conspicuous discolouration in old forestry reports or recovered from systematic field observations. Twenty-two outbreaks are discernible within the period 1800-1983. A peak reduction in larch growth occurred at intervals of 8.86ǃ.01 years in the following years: 1801, 1811, 1820, 1830, 1838, 1846, 1857, 1867, 1875, 1884, 1892, 1901, 1909, 1918, 1926, 1936, 1945, 1953, 1963, 1972, 1980, and 1987. Our objective method based on ring measurements and comparison with non-host trees was compared with qualitative techniques based on the visual detection of conspicuous latewood anomalies. Larch in the Briançonnais (which experiences a continental climate) are infested first, whereas the Tarentaise region exhibits a much weaker impact of LBM. Complete tree recovery was observable 3 years after outbreak peaks.     

Shade tolerance of <Emphasis Type="Italic">Ailanthus altissima</Emphasis> revisited: novel insights from southern Switzerland

The tree of heaven (Ailanthus altissima (Mill.) Swingle) is considered to be an early-successional, gap-obligate pioneer species with vigorous height growth, low shade tolerance, early fecundity and large seed production. It is a highly invasive species in many temperate and Mediterranean ecosystems outside its natural range, especially after disturbance. Due to its low shade tolerance, the potential of A. altissima to colonise undisturbed forests is thought to be low. In this study we analysed the potential of juvenile A. altissima to grow and survive in sweet chestnut (Castanea sativa Mill.) forests in southern Switzerland. We used hemispherical photography to assess the light conditions of 204 individuals of A. altissima (31 % generative, 69 % vegetative) aged between 1 and 7 years (median: 3 years) in six sites. Generative (seed-borne) and vegetative (clonal ramet) offspring of A. altissima are able to grow in light conditions well below the requirements of shade-intolerant tree species such as European larch (Larix decidua Mill.) and Scots pine (Pinus sylvestris L.). The relatively low light conditions found to be sufficient for the growth and survival of generative regeneration of A. altissima suggest a higher shade tolerance for this species than previously stated, at least for early regeneration. Consequently, the colonisation frontier of A. altissima should be intensively monitored in both forest openings but also in closed canopy forests in the vicinity of seed-bearing A. altissima.     

Rates of spontaneous hybridization and hybrid recruitment in co-existing exotic and native mature larch populations

Exotic larch (Larix decidua Mill., Larix kaempferi (Lamb.) Carr. and Larix sibirica Ledeb.) and their hybrids have been introduced into Canada to test adaptation and growth for reforestation purposes. For tree breeding operations, provenance trials and seed orchards of exotic larches and their hybrids have been established in arboreta adjacent to natural forest stands of the indigenous species Larix laricina (Du Roi) K. Koch. This proximity offers an opportunity to measure spontaneous hybridization and the recruitment of hybrids. Using a combination of two chloroplast and three mitochondrial DNA markers, we estimated the rate of spontaneous hybridization taking place between native (L. laricina) and exotic (L. decidua, L. kaempferi, L. sibirica) species of larch in two arboreta. More than 5,000 seeds and 1,000 recruits from the two experimental sites were sampled and genetically identified. The occurrence of spontaneous hybridization between L. laricina and the exotics was found both in seeds and from recruits freely established near the plantations. A low hybridization rate (3 % or less) was observed among the seeds of L. laricina maternal trees. Spontaneous native-exotic hybrids were also observed to establish and survive in the environment under the current climatic conditions at a similarly low rate. However, a much higher and variable rate of establishment was observed for recruits with exotic components (up to 34.6 % at one site). These results indicate that the spontaneous hybridization and the recruitment of hybrid and exotic recruits do occur and should be taken into consideration in the management and establishment of plantations of fast-growing species with exotic components.     

Changes in radial tree growth for Picea abies, Larix decidua, Pinus cembra and Pinus uncinata near the alpine timberline since 1750

Changes in radial growth of the four coniferous species growing in the French Alps near the upper treeline are investigated. Thirty-seven populations of Norway spruce [Picea abies (L.) Karst.], European larch (Larix decidua Mill.), Swiss stone pine (Pinus cembra L.) and mountain pine (Pinus uncinata Mill. ex Mirb.) were sampled by taking 1320 cores and analysing tree-ring widths. Sites were chosen in various climatic conditions (macroclimate and aspect) and on two kinds of bedrock in order to take into account the ecological behaviour of these species. Belledonne, Moyenne-Tarentaise, Haute-Maurienne and Briançonnais areas were sampled along increasing gradients of summer aridity and winter continentality. The calculation of time series after removing the age trend brings strong evidence for an increase in radial growth during the two last centuries, but with different stages and fluctuations for each species. This growth trend is significantly enhanced since 1860 for the spruce, and since 1920 for the two pine species. Furthermore, it also appears on Larix decidua with the same pattern despite periodical growth reduction due to attacks of the larch bud moth (Zeiraphera diniana Gn.). The analysis of ring-widths at a given cambial age reveals that this enhanced phenomenon is observed especially during the tree’s early years (25–75 years). The analysis of four regional climatic series, and three longer series of temperature (in farther single sites) reveals synchronous decadal fluctuations and an evident secular increase in minimum temperatures (especially in January and from July to October), that may be involved in tree-growth enhancement. Thermic amplitudes are significantly reduced during the whole growing period, what is more pronounced in Belledonne, the most oceanic region. Long term growth changes are well described by stepwise regression models, especially for the pine species. These models involved both a linear trend (CO₂ concentration or N-deposition) and low frequency of Turin monthly temperatures. However, they show different patterns than those observed from response functions at a yearly scale.     

Effects of experimental stem burial on radial growth and wood anatomy of pedunculate oak

In dendrogeomorphology, abrupt changes in wood anatomy are frequently used to date the exact year of burial and exposure events. However, few studies have addressed the precision and underlying mechanisms of these changes. In a field experiment, performed in a drift-sand area in the Netherlands, we buried the stems of mature pedunculate oak trees (Quercus robur L.) up to a height of 50 cm and analysed the responses in ring width and earlywood-vessel characteristics, while monitoring the course of temperature above and below the soil surface.After 3 years of stem burial, we found no significant differences in ring width and earlywood-vessel characteristics between control and buried trees both above and below the burial level. Burial however strongly reduced temperature amplitude and the occurrence of sub-zero temperatures around the buried stems. All buried trees formed epitropic roots that grew upward into the new sediment layer, but no adventitious roots were formed on the buried stems. Irrespective of the burial treatments, we found that the mean ring width was largest at the original stem base and lowest at breast height. In contrast, vessel sizes were significantly larger at breast height compared with the stem base. Differences in vessel density barely differed between years and heights.In our field experiment on mature pedunculate oak trees, the burial of stems by 50 cm of drift sand did not induce any local growth suppression or detectable changes in wood anatomy. As wood-anatomical changes in response to burial have previously been reported for trees that had formed adventitious roots, we stress the role of adventitious-root formation as a possible trigger behind the local changes in wood anatomy, reflecting a functional change of a buried stem towards a root. Based on our field experiment, it seems unlikely that years of shallow or moderate burial events (≤50 cm) can be reconstructed using the wood structure of buried stems. As epitropic roots develop quickly after burial, dating such roots may potentially yield better estimates of burial events. Further research on the relation between adventitious root and changes in stem anatomy is needed to ascertain the precision of dating sand-burial events using tree rings.     

Xylem plasticity allows rapid hydraulic adjustment to annual climatic variability

Thanks to acclimation, trees overcome environmental changes and endure for centuries. The anatomy of water conducting cells is an important factor determining plant success. Forming cells are coupled with the environment and their properties are naturally archived in the wood. Its variability across tree rings can thus provide a retrospective of plant’s hydraulic adjustments. In this work, we measured lumen and wall thickness of tracheids along tree-rings to explore how trees regulate their conducting system under variable plant-water conditions. Tracheids were measured along 51 dated rings of five mature Larix decidua and Picea abies trees from a low elevation site. Anatomical-based chronologies of annual growth performance, hydraulic conductance and safety, and construction costs were built. Similarities among chronologies and the relation to monthly climate data were analyzed. Most parameters displayed high annual plasticity which was partly coherent among trees and mostly associated with radial growth. In general, summer drought reduced growth and potential hydraulic conductivity of the forming ring, and increased hydraulic safety and construction costs. To evaluate the functional relevance of the annual acclimation, the conductivity of the forming ring relative to the entire sapwood needs to be assessed.     

Growth trends and dynamics in sub‐alpine forest stands in the Varaita Valley (Piedmont,Italy) and their relationships with human activities and global change

Abstract. A study of the forest lines, tree lines and the structures of the sub‐alpine forest was performed in Vallone Vallanta and in Alevé forest in the Varaita Valley (Cottian Alps, Piedmont, Italy). Forest‐ and tree lines were analysed over 1728 ha while forest structures were studied on six 3000‐m² plots located at the tree line (2), at the forest line (2) and inside the sub‐alpine forest (2). Dendro‐ecological analysis of living plants and stumps showed that Larix decidua was more abundant in the past than today and that Pinus cembra has expanded, both upwards and within sub‐alpine forests. Age structure analysis revealed that the current sub‐alpine forest stands were established 200–220 yr ago, probably following a clearcut. At the forest lines the tree density decreases, and some trees are more than 500 yr old, whereas at the tree lines most of the trees (almost exclusively Pinus cembra) are younger than 100 yr. Growth dynamics were investigated both by observing Basal Area Increment (BAI) in the old and dominant trees, and by comparing the BAIs of classes of trees with a given cambial age range in different time periods. The results showed that the growth rates of mature Pinus cembra and Larix decidua had increased. These increments are more substantial for Pinus than for Larix. The growth rate of young trees (< 100 yr) of both species has decreased over recent decades. This could be due to competition caused by increased tree densities that have resulted from a decrease in grazing.     

Comparing growth phenology of co-occurring deciduous and evergreen conifers exposed to drought

Plant phenological events are influenced by climate factors such as temperature and rainfall. To evaluate phenological responses to water availability in a Spring Heath-Pine wood (Erico-Pinetum typicum), the focus of this study was to determine intra-annual dynamics of apical and lateral growth of co-occurring early successional Larix decidua and Pinus sylvestris and late successional Picea abies exposed to drought. The effect of reduced plant water availability on growth phenology was investigated by conducting a rainfall exclusion experiment. Timing of key phenological dates (onset, maximum rate, end, duration) of growth processes were compared among species at the rain-sheltered and control plot during 2011 and 2012. Shoot and needle elongation were monitored on lateral branches in the canopy at c. 16 m height and radial growth was recorded by automatic dendrometers at c. 1.3 m height of >120 yr old trees. Different sequences in aboveground growth phenology were detected among the three species under the same growing conditions. While onset of radial growth in April through early May was considerably preceded by onset of needle growth in Larix decidua (5–6 weeks) and shoot growth in Pinus sylvestris (c. 3 weeks), it occurred quite simultaneously with onset of shoot growth in Picea abies. Low water availability had a minor impact on onset of aboveground growth, which is related to utilization of stored water, but caused premature cessation of aboveground growth. At the control plot mean growing season length was 130 days in Pinus sylvestris, 95 days in Larix decidua and 73 days in Picea abies, supporting the hypothesis that early successional species are resource expenders, while late successional species are more efficient in utilizing resources and develop safer life strategies. High synchronicity found in culmination of radial growth in late spring (mid-May through early June) prior to occurrence of more favourable environmental conditions in summer might indicate sink competition for carbohydrates to belowground organs. This is supported by completion of apical growth in mid June in all species, except for needle growth of Pinus sylvestris which lasted until early August. Phenological observations of conifers exposed to drought revealed that tree water status early during the growing season determines total annual aboveground growth and, besides temperature, species-specific endogenous and/or environmental factors (most likely photoperiod and/or different threshold temperatures) are involved in controlling apical and lateral growth resumption after winter dormancy.     

Source-driven remobilizations of nutrients within stem wood in Eucalyptus grandis plantations

Nutrient remobilizations in tree ligneous components have been little studied in tropical forests. A complete randomized block design was installed in Brazilian eucalypt plantations to quantify the remobilizations of phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), and sodium (Na) within stem wood. Three treatments were studied: control with neither K nor Na addition (C), 3 kmol ha^?1 K applied (+K), and 3 kmol ha^?1 Na applied (+Na). Biomass and nutrient contents were measured in the stem wood of eight trees destructively sampled at 1, 2, 3 and 4 years after planting in each treatment and annual rings were localized on discs of wood sampled every 3 m in half of the trees. Chemical analyses and wood density measurements were performed individually for each ring per level and per tree sampled. Nutrient remobilizations in annual rings were calculated through mass balance between two successive ages. Our results show that nutrient remobilizations within stem wood were mainly source-driven. Potassium and Na additions largely increased their concentration in the outer rings as well as the amounts remobilized in the first 2 years after the wood formation. The amount of Na remobilized in annual rings was 15 % higher in +Na than in +K the fourth year after planting despite a 34 % higher production of stem wood in +K leading to a much higher nutrient sink. A partial substitution of K by Na in the remobilizations within stem wood might contribute to enhancing Eucalyptus grandis growth in K-depleted soils.     

火烧伤害对兴安落叶松树干径向生长的影响

在大兴安岭塔河林业局瓦拉干林场22支线截取3个兴安落叶松(Larix gmelinii)火疤圆盘,预处理和交叉定年后,测定垂直年轮(径向)和沿年轮(横向)方向2次火烧间的年轮宽度变化.通过生长趋势拟合、火烧后生长面积与正常(理论)生长面积比较,分析火烧伤害对兴安落叶松树干生长的影响.结果表明:火烧后横向年轮生长呈现出窄—宽—正常的趋势,这种趋势可以用3次曲线来拟合,即y=0.01x3-0.14x2+ 0.85x+ 0.56(DF=3,F=227.7,P＜0.0001,y为平均年轮宽度,x为偏离火烧点距离).在本文分析的3个圆盘中,火烧后的横向生长释放都出现在距火烧部位1/3范围内.火烧后径向上年轮宽度变化规律也较为明显,即在火灾后的几年时间内,年轮宽度值较火灾前都呈增加趋势.火烧后3个圆盘的径向生长释放持续时间分别为14、5a和11a.通过面积计算整体来看,火烧导致圆盘1较正常情况下损失了67.2％,圆盘2比正常情况下增加了6.6％,圆盘3增加了13.7％.上述分析表明,火烧导致的径向生长损失,可以较好地由横向和径向生长释放予以补偿,但是具体补偿多少因烧伤程度不同而有差异.火烧可直接改变树干径向生长变化,同时,这种变化也受其他因素影响,例如像树龄、树种特性、火灾间隔期等因素,这些因素的差异就可能导致补偿程度的不同.     

Provenance tests for survival and growth of 50-year-old Japanese larch (Larix kaempferi) trees related to climatic conditions in central Japan

Survival and growth of Japanese larch (Larix kaempferi) from 25 provenances in the natural distributional range were examined at three test sites in central Japan. Climatic conditions of the sites and provenances indicated a contrast between northwest and southeast locations across the distributional range. Density, height, diameter, and crookedness of stems and width and length of branches were measured for 50-year-old trees, and per-area cross-sectional area and volume of stems were calculated. Variance components among the provenances were large in the stem area and volume (27–33 %) and the branch width (26 %); those among the sites were large in the stem height, diameter, and density (70–81 %); and those of interaction between sites and provenances were small (0–12 %) in all the variables. Principal component analyses showed that eastern provenances exhibited large size of stems and branches at most sites. Growth rate and maximum size at each site were estimated from fitting the stem volume at three ages (12, 31 or 32, and 50 years) to growth curves in logistic functions. Eastern provenances, where seasonal variation is least in temperature and most in precipitation, showed the highest performance of stem and branch growth. The findings from our long-term survey of the L. kaempferi provenance tests suggest climatic effects on survival and growth traits and potential provenances from which to select new breeding materials.     

Expanding forests and changing growth forms of Siberian larch at the Polar Urals treeline during the 20th century

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.     

Responses of young trees to wind and shading: effects on root architecture

Two wind tunnels were designed to detect influences of windon the development of the root systems of young trees: Piceasitchensis and Larix decidua were grown in the first and L.decidua only in the second. In the second experiment, the taproot of each L. decidua seedling was removed in order to mimicthe formation of a shallow root-plate. Responses of shoot growthto wind stress were small, although uneven irradiance levelsresulted in asymmetric growth of both shoots and roots; thedistribution of root biomass around the tree was related spatiallyto that of shoot growth In both experiments there was an increase in the number of largewindward and leeward roots in both species. In the first experiment,the sum of the crosssectional area (     

Couplings in cell differentiation kinetics mitigate air temperature influence on conifer wood anatomy

Conifer trees possess a typical anatomical tree‐ring structure characterized by a transition from large and thin‐walled earlywood tracheids to narrow and thick‐walled latewood tracheids. However, little is known on how this characteristic structure is maintained across contrasting environmental conditions, due to its crucial role to ensure sap ascent and mechanical support. In this study, we monitored weekly wood cell formation for up to 7 years in two temperate conifer species (i.e., Picea abies (L.) Karst and Larix decidua Mill.) across an 8°C thermal gradient from 800 to 2,200 m a.s.l. in central Europe to investigate the impact of air temperature on rate and duration of wood cell formation. Results indicated that towards colder sites, forming tracheids compensate a decreased rate of differentiation (cell enlarging and wall thickening) by an extended duration, except for the last cells of the latewood in the wall‐thickening phase. This compensation allows conifer trees to mitigate the influence of air temperature on the final tree‐ring structure, with important implications for the functioning and resilience of the xylem to varying environmental conditions. The disappearing compensation in the thickening latewood cells might also explain the higher climatic sensitivity usually found in maximum latewood density.     

3D analysis of anatomical reactions in conifers after mechanical wounding: first qualitative insights from X-ray computed tomography

The ability of trees to recover from damage beyond the last-formed periderm as well as the drivers and nature of associated wound reactions have been studied for more than two centuries using macroscopic (desiccation, aeration or discoloration of wood) and microscopic approaches (anatomical and chemical reactions). However, no studies currently exist which address large-scale macroscopic and microscopic reactions surrounding wounds in the tangential, axial, and radial directions over continuous segments of tree stems. This note explores the potential of 3D X-ray computed tomography in assessing effects of wounding under natural conditions in European conifers (Abies alba, Larix decidua, Picea abies). We present results from a pilot study and qualitatively evaluate the potential of the approach used in assessing and illustrating the formation and spread of de-differentiated xylem parenchyma cells, xylem decay compartmentalization, resin ducts, and stabilizing compression wood cells.     

The influence of stem guying on radial growth,stem form and internal resin features in radiata pine

Key message

Stem guying to prevent wind-induced swaying of radiata pine trees resulted in significant changes in radial growth, but did not affect the frequency of compression wood or resin features.

Abstract

Mechanical stress resulting from wind forces acting on trees can cause a number of direct and indirect effects ranging from microscopic changes in cambial activity through to stem breakage and uprooting. To better understand these effects on radial stem growth and wood properties, an experiment was established in a 13-year-old radiata pine (Pinus radiata D Don) stand in which 20 trees were guyed to prevent them from swaying. Radial growth was monitored in these trees and 20 matched controls at monthly intervals for 5 years. The trees were then felled and radial growth, resin features and compression wood were assessed on cross-sectional discs taken at fixed locations up the stem. There was a significant reduction in radial growth at breast height (1.4 m above the ground) in the guyed trees, but an increase in growth immediately above the guying point. A total of 277 resin features were observed in the growth rings formed following guying. The overall frequency of such features was related to height within the stem and annual ring number. No effect of stem guying was found on the incidence of compression wood. Interestingly, the distribution of resin features also did not differ between guyed and un-guyed trees. There was no evidence of a link between stem restraint as a result of guying and the incidence of resin features, suggesting that other factors, such as soil moisture may be more influential.     

Disturbance dynamics and history of an old-growth subalpine fir (Abies fargesii) forest in central China

Disturbance history of an old-growth subalpine fir (Abies fargesii) forest in the Shennongjia Mountains of central China was reconstructed using dendroecological methods. Increment cores were extracted from 468 trees within six 100 m × 50 m permanent transects distributed across the old-growth subalpine fir forest of 300 ha. Growth patterns of 299 fir cores were examined for abrupt increases in radial growth to indicate formation of past canopy gaps and for rapid early radial growth to indicate establishment in past canopy gaps. The results showed that 70.8 % of the canopy fir trees experienced an average of 0.78 (ranging from 0 to 2) major release event for an average of 15.8 (ranging from 10 to 24) years, and an average of 1.94 (ranging from 0 to 3) moderate release events for an average of 25.6 (ranging from 10 to 36) years before they reached canopy. Recruitment pulse of trees coincided temporally with the peak of disturbance rate from the 1900s to the 1910s, suggesting occurrence of intense disturbance events during the time period. Radial growth analyses indicated that a history with small-scale disturbance events has resulted in the formation of the old-growth subalpine fir forest, and stand-replacing disturbances might not be necessary for the development of the forest. This study provides strong evidence that there are substantial variations in the disturbance severity and frequency over time. Most disturbance events might rather cause treefall gaps than clear large areas of forest at once. Thus, the old-growth subalpine fir forest experienced frequent gap-scale disturbances and few large-scale disturbances in its development history.     

Tree growth and competition in a Betula platyphylla-Larix cajanderi post-fire forest in central Kamchatka

BACKGROUND AND AIMS: Fire is the dominant disturbance in central Kamchatka boreal forests, yet patterns and mechanisms of stand recovery have not been investigated. METHODS: Measurements were made of 1433 stems > or =1.3 m height and annual radial increments of 225 randomly selected trees in a 0.4-ha plot of a 53-year-old fire-origin mixed-species stand to examine the spatio-temporal variation in establishment, growth, size inequality and the mode of competition among individual trees. Growth variations were related to tree size, age and local interference with neighbours. KEY RESULTS: Betula platyphylla formed the main canopy following a fire in 1947, with Larix cajanderi and Pinus pumila progressively reinvading the lower tree and shrub stratum. Most B. platyphylla originated from sprouts in small patches (polycormons) during the first 15 post-fire years. Betula platyphylla had normal distributions of diameter and age classes, but negatively skewed height distribution, as expected from shade-intolerant, pioneer species. Larix cajanderi had fewer tall and many short individuals. The smaller and younger B. platyphylla grew disproportionately more in diameter than larger trees from 1950 to 1975, and hence stem size inequalities decreased. The reverse trend was observed from 1995 to 2000: larger trees grew more, indicating an increasing asymmetry of competition for light. Betula platyphylla had steady diameter growth in the first 25 post-fire years, after which the growth declined in smaller trees. Neighbourhood analysis showed that the decline resulted from increased competition from taller neighbours. CONCLUSIONS: The observed growth patterns suggest that mode of interactions altered during stand development from early stages of weak competition for soil resources released by fire to later stages of asymmetric competition for light. Asymmetric crown competition started later than reported in other studies, which can be attributed to the lower stem density leaving much space for individual growth, greater relative importance of below-ground competition in this site of nutrient-poor volcanic soil, and the vegetative origin of B. platyphylla. Larix cajanderi growing under B. platyphylla had steady diameter growth during the first 20 years, after which growth declined. It is suggested that early succession fits the tolerance model of succession, while inhibition dominates in later stages.     

Spatiotemporal distribution of growth releases and suppressions along a landslide body

Our study analyzes the growth response (release or suppression) of Norway spruce trees growing along a landslide zone in eastern Czech Republic. A total of one hundred and eighty-six increment cores were extracted from Norway spruce (Picea abies (L.) Karst.) individuals, which were affected by two different disturbances, the Girová landslide in May 2010 and an anthropogenic cut-off in the neighborhood of the landslide later the same year. Growth changes were analysed in three zones that were defined according to the type of effect the disturbance had on the surrounding vegetation. The aims of our study are to (i) detect growth changes in trees that survived the disturbances in 2010, (ii) evaluate the delay time of the growth reaction to the disturbances, (iii) compare how growth changes differed in response to the two different disturbance types (natural landslide vs. anthropogenic harvesting and deforestation), and (iv) investigate spatio-temporal differences in the growth changes. Our results indicate that tree growth changed in response to the altered environmental conditions following the 2010 disturbances. The changes vary depending on the intensity of the disturbance and its effect on the forest stand. Trees in Zone 1 responded with a slight growth release (14.6 % of responding trees), which was strongest after 2013. However, growth suppression (85.4 %) was the dominant reaction in Zone 1; with the majority of trees showing growth suppression in 2014, four years after the events. The strongest and most abrupt growth release (66.1 %) occurred in Zone 2 in 2011, after a one-year delay. Since 2010, we have observed a trend of growth suppression (33.9 %) in this zone. In Zone 3, following a one-year delay, growth release has occurred gradually in about a third of the trees (37.0 %). The majority of trees in this zone have responded with growth suppression (63.0 %) in 2010 and with a three-year delay after 2013. We also found differences in how tree growth responds to anthropogenic and landslide disturbances, as well as spatio-temporal differences related to the extent of post-disturbance changes.     

Slow lifelong growth predisposes Populus tremuloides trees to mortality

Widespread dieback of aspen forests, sometimes called sudden aspen decline, has been observed throughout much of western North America, with the highest mortality rates in the southwestern United States. Recent aspen mortality has been linked to drought stress and elevated temperatures characteristic of conditions expected under climate change, but the role of individual aspen tree growth patterns in contributing to recent tree mortality is less well known. We used tree-ring data to investigate the relationship between an individual aspen tree’s lifetime growth patterns and mortality. Surviving aspen trees had consistently higher average growth rates for at least 100 years than dead trees. Contrary to observations from late successional species, slow initial growth rates were not associated with a longer lifespan in aspen. Aspen trees that died had slower lifetime growth and slower growth at various stages of their lives than those that survived. Differences in average diameter growth between live and dead trees were significant (α = 0.05) across all time periods tested. Our best logistical model of aspen mortality indicates that younger aspen trees with lower recent growth rates and higher frequencies of abrupt growth declines had an increased risk of mortality. Our findings highlight the need for species-specific mortality functions in forest succession models. Size-dependent mortality functions suitable for late successional species may not be appropriate for species with different life history strategies. For some early successional species, like aspen, slow growth at various stages of the tree’s life is associated with increased mortality risk.