Stem Diameter Growth of Scots Pine Trees after Increased Mechanical Load in the Crown during Dormancy and (or) Growth

A field experiment with a 2 x2 factorial block design (W_xS_x)was conducted in northern Sweden where the mechanical loadsin the crowns of sixteen 2.5m high Scots pine (Pinus sylvestrisL.)trees were increased during one winter (W₁, dormant period)and (or) summer (S₁, growth period). Trees treated were loadedwith five 2kg bags hung over mid-crown branches close to thestem, i.e. 10kg per tree. After treatment, all trees were leftto grow untreated for one additional year. Trees were then cutat ground level and annual ring widths for the last 5 yearswere measured on stem discs taken at 1, 5, 10, 15, 20, 30 and50% of tree height. Differences between treatments were analysedwith two-way factorial ANOVA. Accumulated treatment responsewas positive for winter loading (W₁S_x) at all levels, and statisticallysignificant at 1, 15 and 20% of tree height. Summer loading(W_xS₁) had positive effects at the lowest and middle parts ofthe stem, and negative in between. No statistically significanttwo-way interaction (W xS) was observed. Results support thehypothesis that Scots pine trees can retain information aboutmechanical forces acting on their stems during winter, and respondto this during the following growth period. The results alsosuggest that stem form of trees in boreal forests may be stronglyaffected by winter conditions. Stem form; mechanical perturbation; Scots pine; Pinus sylvestris; dendrometer; diameter; growth; dormancy; thigmomorphogenesis; wind; sway     

Root biomass and rooting profile of naturally regenerated beech in mid-elevation Scots pine woodlands

The natural establishment of shade-tolerant forest species such as beech (Fagus sylvatica L.) occurs in naturally regenerated Scots pine (Pinus sylvestris L.) woodlands that develop on former pastures and cultivated lands. To examine possible effects of underground competition in beech establishment, we studied the root biomass and the rooting profile of 53 mixed Scots pine-natural beech woodlands in French mid-elevation volcanic areas. Stands were arranged along a maturation gradient. Roots were sampled using the root-auger technique (0 to 75 cm depth every 15 cm). In addition, 23 young beeches were uprooted to study the entire root system. Total beech fine-root biomass was closely correlated with most beech aerial characteristics, (e.g., height, diameter and girth), and correlated moderately with tree age. However, it correlated poorly with basic competition indices such as stand density and basal area. Conversely, competition indices including vertical dimensions of competing trees were correlated with the underground biomass, probably as a result of redundancy with beech height. The rooting profile (fine roots, < 5 mm) of beech and pine were quite similar, and did not change significantly along the stand maturation gradient. Beech has a heart-shaped root system while pine is more plate-like and dimorphic. Beech fine-root biomass progressively surpassed pine biomass throughout the soil layers, thus confirming that it is dynamic and competitive in mature mixed stands. The coexistence of the root systems of beech and pine in the same soil layers presumably results in strong underground competition.     

Rooting strategy of naturally regenerated beech in Silver birch and Scots pine woodlands

This study investigated the belowground development and strategy of late-successional European beech (Fagus sylvatica L.) in ageing natural Scots pine (Pinus sylvestris L.) and Silver birch (Betula pendula Roth.) woodlands in a French volcanic mid-elevation area. For this purpose root biomass, root profile and fine-root architecture of competitor trees were examined in 53 mixed pine–beech and 42 birch–beech woodlands along a stand maturation gradient, using the root auger technique (0–75-cm). The total beech fine-root biomass highly correlated with aerial dimensions such as stem height and girth, whereas it moderately correlated with its age, thus indicating the effects of competition. Basic stand biometric data such as stand density and basal area had no significant effect on beech root biomass. Conversely, competition indices taking into account the vertical dimensions of competitor trees were efficient, probably due to redundancy with beech height. At similar age and height, beeches under birch had a greater belowground development than beeches under pine. Each species exhibited specific rooting pattern and plasticity of fine-root architecture along the gradients of stand maturation and competition. Beech had a heart-shaped rooting habit in both mixings, which strongly increased along stand maturation. Its fine-root system adopted a foraging strategy to respond to increasing stand competition. The Scots pine fine-root system was plate-like and showed a low morphological plasticity, thus presumably a conservative strategy. Silver birch exhibited a high biomass and a foraging capacity in the topsoil but a loose root system in the subsoil. The coexistence of pine and beech roots in the upper soil presumably leads to a high belowground competition. Beech root system becomes predominant throughout the soil profile and it adopts an efficient foraging strategy, but at the expense of its belowground development. Conversely, the niche partitioning strategy between beech and birch may explain why beech develops strongly belowground in spite of the fact that birch has a dense rooting and a competitive fine-root architecture. As a consequence, beech mid-term regeneration and development may be facilitated under birch as compared with pine.     

Crown structure and leaf area of the understorey species Prunus serotina

A detailed biometrical study of the exotic understorey invader Prunus serotina (Ehrh.) was carried out in a mixed coniferous forest stand in northern Belgium. Based on detailed destructive measurements of eight selected model trees, allometric relations of tree height, crown projected area, woody and leaf dry mass and leaf area on tree diameter at breast height (DBH) were derived. The scaling-up procedure from the tree to the stand level was done using the frequency distribution of DBH obtained at the selected experimental plot. The vertical and radial distributions of the tree foliage were estimated by the “cloud” technique. The vertical profile of leaf area showed a bimodal distribution pattern with maxima at heights of 4 and 6 m above the ground. The leaf area index (LAI) of the understorey Prunus serotina as estimated by the described up-scaling procedure (5.1) was significantly higher than the LAI (2.6) as measured by a plant canopy analyser and was also significantly higher than the LAI of the overstorey species Scots pine (1.5–3.0). The LAI of a neighbouring Rhododendron understorey reached only 1.25. This study emphasises the importance of an exotic understorey species in the total leaf area of mixed coniferous forests which might have important implications for the energy and mass exchanges of the entire forest.     

Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest

Summary The spatial distribution of seedlings of the dominant perennial plant species (Pinus halepensis, Cistus salviifolius, Rhus coriaria) and may annual species was studied after a wild fire in an eastern Mediterranean pine forest. The spatial distribution of all seedlings is affected by the location of the old burned pine trees. Seedling density of Pinus and Cistus is higher at a distance from the burned pine canopy and lower near the burned pine trunk. It is also higher beneath small burned pine trees than under big ones. Rhus seedling density is higher under big burned pine trees and also near the burned trunks. Seedlings of Pinus, Cistus and Rhus growing under the burned canopy of big pine trees tend to be taller than seedlings under small ones or outside the burned canopy. Most annual species germinate and establish themselves outside the burned canopies, and only a few annual species are found beneath them. It is suggested that variation in the heat of the fire, in the amount of ash between burned pine trees of different sizes, and in the distance from the burned canopy are responsible for the observed pattern of seedling distribution. The possible ecological significance of the spatial pattern of seedlings distribution and their differential growth rate are discussed.     

Recovery of vegetation in a Caledonian pinewood after fire

Summary

Following a wildfire in the Caledonian pinewood at Shieldaig, Wester Ross in late March 1974, the recovery of the vegetation, particularly Scots pine (Pinus sylvestris) seedlings, was monitored in twenty sample plots. Almost all trees under 5 cm diameter at breast height were immediately killed by the fire and of the larger trees, 45 per cent of the Scots pine and 60 per cent of the birch (Betula pendula) had died by 1980. Regeneration of Scots pine had occurred to the extent of about 2,500 seedlings ha^-1 by 1980. Recovery of the field layer species varied; species with rhizomes grew up quickly but the re-establishment of cover by Calluna vulgaris and many bryophytes was slow.     

Growth allocation between height and stem diameter in nonsuppressed reproducing Abies mariesii trees

To understand the growth patterns with respect to competition and leaf‐mass increase in reproducing trees, growth allocation between height and stem diameter was examined for nonsuppressed reproducing Abies mariesii trees in a subalpine forest in northern Honshu, Japan. The growth allocation was analyzed by dividing the relative growth rate of the stem volume into the relative contributions of height and stem‐diameter growth. During a 9‐yr period, height growth and seed‐cone production showed obvious annual variation, while stem‐diameter growth recorded moderate variation. For two of three years of seed‐cone production during the 9‐yr period, trees with larger seed‐cone production were associated with less height growth in the following year of seed‐cone production; however, there was no trend of height growth in the year of seed‐cone production. In the following year of mast seeding, trees with larger stem‐volume growth were associated with less height growth. This trend was also shown for the relationship between the cumulative stem‐volume growth during the 9‐yr period and growth allocation to height, suggesting that trees with a larger biomass increase depress the allocation of photosynthate to competition with a large expenditure for reproduction. In contrast to this, trees with a smaller biomass increase might allocate photosynthate to competition with surrounding trees. The results of this study suggest that an increase in reproductive organs during life history and annual variation in reproduction are closely associated with the growth patterns of the stem in A. mariesii trees.     

Heartwood and sapwood development within maritime pine (<Emphasis Type="Italic">Pinus pinaster </Emphasis>Ait.) stems

Heartwood and sapwood development in maritime pine (Pinus pinaster Ait.) is reported based on 35 trees randomly sampled in four sites in Portugal. It was possible to model the number of heartwood rings with cambial age. The heartwood initiation age was estimated to be 13 years and the rate of sapwood transformation into heartwood was 0.5 and 0.7 rings year^–1 for ages below and above 55 years, respectively. Reconstruction of heartwood volume inside the tree stem was made by visual identification by image analysis in longitudinal boards along the sawn surfaces. This volume was integrated into the 3D models of logs and stems developed for this species representing the external shape and internal knots. Heartwood either follows the stem profile or shows a maximum value at 3.8 m in height, on average, while sapwood width is greater at the stem base and after 3 m remains almost constant up the stem. Up to 50% of tree height heartwood represents 17% of stem volume, in 83-year-old trees and 12–13% in 42 to 55-year-old trees. Tree variables such as stem diameter, DBH and tree total height were found to correlate significantly with the heartwood content.     

Relationship between growth rates and xylem hydraulic characteristics in young, mature and old-growth ponderosa pine trees

The first objective of the present study was to quantify the effects of tree age and stem position on specific conductivity (k_s), vulnerability to embolism and water storage capacity (capacitance) in trunks of young, mature and old‐growth ponderosa pine. The second objective was to determine relationships between hydraulic characteristics and radial and height growth rates to increase the understanding of possible tradeoffs. Within sapwood at all heights and in all ages of trees, outer sapwood had 25–60% higher k_s than inner sapwood. The water potential at which embolism started (air entry point) was 1.3 MPa lower in inner sapwood than outer sapwood within the mature trees, but there was no difference in the other trees. There was no significant difference in capacitances between the tops of the old growth trees, the mature trees and the young trees. Taking all data together, the capacitances increased sharply with an increase in k_s and an increase in vulnerability to embolism. The hydraulic characteristics of the three age classes were correlated with the height growth rate but not with the diameter growth rate. Within these age classes, high k_s was associated with the slowest yearly increase in sapwood area and with a low percentage of latewood, whereas high vulnerability to embolism and high capacitance were more closely associated with high height growth rates.     

Development processes and growth pattern of Pinus densiflora stands in central eastern Korea

Stand growth and developmental processes were investigated in Pinus densiflora Siebold et Zucc. stands of different ages in the central eastern region of Korea. Stands were inventoried and five trees per stand were sampled for stem analysis, age estimation, and growth analysis. More than 80% of sampled trees in a stand were established within 3–5 years, and most stands had a single cohort structure. The initial growth of pine seedlings was slow, but the height growth accelerated beyond 2–3 m height, 5–10 years after establishment. Linear growth was maintained until 10–12 m height, at which suppressed trees fell behind and might die out. The young stand was composed of pure pines, while few pine seedlings and saplings were found in the understory of older stands. The peak of diameter growth rate occurred around 5–15 years after tree establishment, implying that competition begins during that period. The pine stand development follows four stages: (1) the young stage when the growth rate increases and peaks; (2) the height competition stage when trees focus on height growth for light while maintaining a narrow DBH and height distribution; (3) the differentiation stage when suppressed trees die out, and the DBH distribution becomes wider; and (4) the mature stage when stands have a multi-canopy structure with a wide DBH and height distribution, while the understory is dominated by other tree species. The changes in growth rates and stand structure through forest development would be implemented to predict alterations of above-ground carbon sequestration rates.     

Connections between climatic variables and the growth and needle dynamics of Scots pine (Pinus sylvestris L.) in Estonia and Lapland

The relationships between climatic variables and Scots pine (Pinus sylvestris L.) growth and needle dynamics were studied in three stands in Estonia and in four stands located near the northern timberline in Lapland. The trees sampled in Estonia had low correlations with the analysed climatic variables (air temperature, precipitation and indices of atmospheric circulation). Moreover, the weak cross-correlation of the time-series of the Estonian sample trees indicated that Scots pine is affected mainly by local factors in that region. In Lapland, however, height increment and needle production correlated strongly among trees within a stand (mean r=0.45 and 0.46, respectively) and between stands (r=0.32 and 0.37). Radial increment also showed a high inter-correlation among the trees within a stand in Lapland (r=0.45). Both height increment and needle production were strongly influenced by the temperature regime of the previous summer in Lapland (mean r=0.64 and 0.64, respectively). Radial increment was correlated with the mean July temperature of the current year (mean r=0.29). The correlations between the indices of atmospheric circulation and tree attributes were weak, while the strongest correlation was between the Ponta Delgada NAO index (PD–NAO) and height increment and needle production in Lapland. Height increment, needle production and radial increment have increased since the 1990s in the trees growing in Lapland. This may indicate a positive effect of climate warming on tree growth in Lapland. In Estonia, where climatic conditions do not limit tree growth, the climate warming seems not to directly influence the growth and needle dynamics of Scots pine.     

Host resistance in defoliated pine: effects of single and mass inoculations using bark beetle-associated blue-stain fungi

1 In 1996, 7000 ha of pine forests were defoliated by the pine looper Bupalus piniaria in south‐western Sweden. 2 The susceptibility of trees of different defoliation classes (0, 30, 60, 90 and 100% defoliation) to beetle‐vectored blue‐stain fungi was tested in inoculation experiments. Forty and 120‐year‐old Scots pine trees were inoculated with ‘single’, i.e. a few inoculations of Leptographium wingfieldii and Ophiostoma minus, two blue‐stain fungi associated with the pine shoot beetle Tomicus piniperda. The young trees were also ‘mass’ inoculated with L. wingfieldii at a density of 400 inoculation points per m² over a 60 cm stem belt. 3 Host tree symptoms indicated that only trees with 90–100% defoliation were susceptible to the mass inoculation. 4 Single inoculations did not result in any consistent differences in fungal performance between trees of different defoliation classes, regardless of inoculated species or tree age class. 5 Leptographium wingfieldii produced larger reaction zones than O. minus, and both species produced larger lesions in old than in young trees. 6 As beetle‐induced tree mortality in the study area occurred only in totally defoliated stands, mass inoculations seem to mimic beetle‐attacks fairly well, and thus seem to be a useful tool for assessing host resistance. 7 As even severely defoliated pine trees were quite resistant, host defence reactions in Scots pine seem to be less dependent on carbon allocation than predicted by carbon‐based defence hypotheses.     

The effect of shoot growth stage of Pinus contorta and Pinus sylvestris on the growth and survival of Panolis flammea larvae

Summary Young Panolis flammea (Lepidoptera: Noctuidae) larvae were unable to survive on the mature (one-year old) needles of Pinus contorta and Pinus sylvestris. Larval growth and survival on the current year's shoots of the 2 pine host plants first incresed, and then decreased as pine shoot development progressed, the effect on survival being more marked at 10° C that 18° C. The survival of larvae on lodgepole pine on field-grown plants transferred to 10° C rose from 26% in March to 87% in mid May, and rose from 3% to 82% on Scots pine in the same period. Larval survival exceeded 70% for about 8 weeks on lodgepole pine and about 6 weeks on Scots pine, this period starting and ending earlier on lodgepole than on Scots pine. The nitrogen, water and phosphorus contents of both pines were at a maximum in May and decline gradually thereafter; the soluble tannin content showed a more complex pattern. These results are discussed in relation to the possible importance of phenological coincidence on the population dynamics of P. flammea with emphasis on its greater abundance on lodgepole pine in Scotland.     

Structural attributes,tree-ring growth and climate sensitivity of Pinus nigra Arn. at high altitude: common patterns of a possible treeline shift in the central Apennines (Italy)

European black pine (Pinus nigra ssp. nigra Arnold) encroachment at increasing elevation has been analyzed at four treeline ecotones of the central Apennines (Italy). The study sites are located along a North-South gradient of 170 km across Marche and Abruzzo regions in Central Italy. The aims of this study were: (i) to detect possible common patterns of structural attributes of black pine regeneration at the treeline ecotone; (ii) to date the seedlings germination and (iii) to assess the climate influence on the pine upward encroachment process also using intra-annual density fluctuations (IADFs) in tree-rings. We sampled 658 encroached black pine trees above the current treeline to the mountain top. All individuals were mapped and their basal stem diameter, total height, annual height increments and other structural attributes measured. One increment core was extracted from stem base of most samples for cambial age determination and detection of intra-annual density fluctuations (IADF). At two sites we also extracted cores at DBH from forest trees to assess climate–growth relationships of black pine. We used multivariate analysis (PCA) to explore the correlation structure of the main tree attributes, regression analysis to relate radial and height increment and dendroclimatic analysis to assess the influence of climate on tree growth and IADF formation.Most black pine trees were located at high altitude and their structural attributes were similar at the four sites where the pine encroachment process started between 30 and 40 years ago featuring similar germination peaks and growth patterns. Black pine is particularly sensitive to maximum temperatures and IADF occurred in mid-late summer with highest frequency peaks between 2003 and 2004. The pine encroachment process, besides the differences of environmental features and land use histories of the four study sites, appears synchronic and spatially diffused. Consistent tree-growth dynamics and the species adaptation to a warming climate are signals envisaging a possible treeline upward shift.     

Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) wood properties in an alkaline air pollution environment

This paper examines the long-term influence on pine (Pinus sylvestris L.) wood properties of alkaline dust pollution (pH 12.3–12.7) emitted over 135 years from a cement plant in Estonia. A study of stemwood physical and mechanical properties in 70–80-year-old Scots pines growing in three zones of different air pollution levels showed serious deviations in comparison with a relatively healthy forest in an unpolluted area. Specimens from polluted trees evidenced smaller sapwood annual ring widths than those from the control trees. At the same time, the number of growth rings in sapwood at breast height increased under pollution. In the polluted areas, percentage of latewood in the annual ring widths was higher than in the unpolluted area. Small amounts of cement dust, which contains elements essential for the mineral nutrition of the trees, might have acted as fertilizer. Pine wood in the polluted stands exhibited increased density, bending strength across the grain, compression strength along the grain and, in some instances, hardness along the grain.     

Fast response of Scots pine to improved water availability reflected in tree‐ring width and δ13C

Drought‐induced forest decline, like the Scots pine mortality in inner‐Alpine valleys, will gain in importance as the frequency and severity of drought events are expected to increase. To understand how chronic drought affects tree growth and tree‐ring δ¹³C values, we studied mature Scots pine in an irrigation experiment in an inner‐Alpine valley. Tree growth and isotope analyses were carried out at the annual and seasonal scale. At the seasonal scale, maximum δ¹³C values were measured after the hottest and driest period of the year, and were associated with decreasing growth rates. Inter‐annual δ¹³C values in early‐ and latewood showed a strong correlation with annual climatic conditions and an immediate decrease as a response to irrigation. This indicates a tight coupling between wood formation and the freshly produced assimilates for trees exposed to chronic drought. This rapid appearance of the isotopic signal is a strong indication for an immediate and direct transfer of newly synthesized assimilates for biomass production. The fast appearance and the distinct isotopic signal suggest a low availability of old stored carbohydrates. If this was a sign for C‐storage depletion, an increasing mortality could be expected when stressors increase the need for carbohydrate for defence, repair or regeneration.     

Relative Importance of Environmental Stress and Herbivory in Reducing Litter Fall in a Semiarid Woodland

We examined the impact of soil stress (low water and nutrient availabilities) and two keystone insect herbivores on pinyon pine (Pinus edulis) needle litterfall. We compared trees growing on two distinct soil types: volcanic cinders, which exhibit pronounced water and nutrient limitation, and sandy-loam soils, which have higher water-storage capacity and nutrient availability. Using two long-term herbivore removal experiments (15 and 18 years, respectively), we also examined the effects of the pinyon needle scale (Matsucoccus acalyptus, which attacks juvenile trees) and the stem-boring moth (Dioryctria albovittella, which attacks mature trees) on pinyon litterfall. These herbivores reach high densities on cinder soils but are absent or occur at much lower levels on sandy-loam soils. Four years of litterfall measurements showed four major patterns. First, independent of herbivory, needle litterfall was 20% lower under trees on high-stress cinder soils than on sandy-loam soils. Second, in agreement with the negative impact of scales on tree growth (that is, a 30% decline in stem growth), trees with scale infestations had 25% lower litterfall rates than trees resistant to scale; however, 15 years of scale-insect removal did not significantly increase needle litterfall. This implies possible intrinsic differences in litter production between scale-resistant and scale-susceptible trees. Third, in contrast with significant negative effects of moth herbivory on tree growth (that is, a 27% decline in stem growth), moth herbivory had no effect on needle litterfall. This, along with increased stem density in moth-susceptible trees, may be evidence of compensatory production. Fourth, there were strong year by soil type and year by scale herbivory interactions, such that in some years the effect on litterfall can be obscured or reversed by some other factor. In summary, soil stress has a strong and predictable effect on needle litterfall, whereas the relationship between insect herbivory and needle litterfall is weaker and depends on the individual herbivore. These effects, however, are mediated by other environmental factors that have considerable annual variation.     

Plasticity in gas‐exchange physiology of mature Scots pine and European larch drive short‐ and long‐term adjustments to changes in water availability

Adjustment mechanisms of trees to changes in soil‐water availability over long periods are poorly understood, but crucial to improve estimates of forest development in a changing climate. We compared mature trees of Scots pine (Pinus sylvestris) and European larch (Larix decidua) growing along water‐permeable channels (irrigated) and under natural conditions (control) at three sites in inner‐Alpine dry valleys. At two sites, the irrigation had been stopped in the 1980s. We combined measurements of basal area increment (BAI), tree height and gas‐exchange physiology (Δ¹³C) for the period 1970–2009. At one site, the Δ¹³C of irrigated pine trees was higher than that of the control in all years, while at the other sites, it differed in pine and larch only in years with dry climatic conditions. During the first decade after the sudden change in water availability, the BAI and Δ¹³C of originally irrigated pine and larch trees decreased instantly, but subsequently reached higher levels than those of the control by 2009 (15 years afterwards). We found a high plasticity in the gas‐exchange physiology of pine and larch and site‐specific responses to changes in water availability. Our study highlights the ability of trees to adjust to new conditions, thus showing high resilience.     

Daily dynamics in xylem cell radial growth of Scots pine (Pinus sylvestris L.)

Daily dynamics of radial cell expansion during wood formation within the stems of 25-year-old Scots pine trees (Pinus sylvestris L.), growing in field conditions, were studied. The samples of forming wood layers were extracted 4 times per day for 3 days. Possible variations in the growth on different sides of the stem, duration of cell development in radial cell expansion phase and dynamics of cell growth in this phase were taken into account. The perimeters of tracheid cross-sections as a reflection of primary cell wall growth were the criterion of growth in a radial direction. For the evaluation of growing cell perimeters a special system for digital processing and image analysis of tracheid cross-sections of the forming wood was used. Growth rate for certain time intervals was estimated by the change in the relation of the perimeter of each observed cell in each of ten tracheid rows in each of 12 trees to the perimeter of the xylem cell of the same row before the expansion. Temporal differences in average values of the relations were estimated by Analyses of Variance. The existence of daily dynamics of Scots pine xylem cell radial growth has been proved. Intensive growth of pine tracheids has been shown to occur at any time of the day and to depend on the temperature regime of the day and the night as well as water supply of stem tissues. Moreover, reliable differences (P = 0.95) in the increment of cell walls during tracheid radial expansion have been found. Pulsing changes of the water potentials both of the cell and the apoplast, as the reason for the fluctuations of radial cell growth rate, were discussed.     

Production of Seeds and Cones and Consequences for Wood Radial Increment in Norway Spruce (Picea Abies (L.) Karst.)

Abstract

The presence of a trade–off between growth and reproduction was tested in four sites in a subalpine Norway spruce (Picea abies (L.) Karst.) forest by measuring annual stem diameter increments at breast height and seed and cone productions during the periods 1962–1985 and 1983–1990, respectively. Trees growing in forest stands near the timber line (about 1900 m above sea level) had the greatest reduction in annual stem diameter increment during mast years; while trees growing at about 1300–1500 m above sea level did not show any reduction. Trees growing at about 1700 m showed only a limited reduction. At the same elevation, trees growing within closed forest stands suffered a greater reduction in stem growth when compared with trees growing at the edge of a cutting.