Gap characteristics and gap regeneration in subalpine old-growth coniferous forests,central Japan

Gap characteristics and gap regeneration were studied in three old-growth stands of subalpine coniferous forests in the northern Yatsugatake and the northern Akaishi mountains, central Japan. With the results of the present study and those of a previous study conducted in another locality, general features of gap characteristics and gap regeneration behavior of major tree species in subalpine coniferous forests of central Japan were summarized and discussed. Of the total 237 gaps investigated in the 14.48 ha of forested area, the percentage gap area to surveyed area, gap density and mean gap size were 7.3%, 17.2 ha⁻¹, and 43.3 m², respectively. The gap size distributions were similar among stands and showed a strong positive skewness with a few large and many small gaps; gaps <40m² were most frequent and those >200 m² were rare. Gaps due to the death of multiple canopy trees comprised 44.7% of the total ones. Canopy trees died in various states; standing dead (42.6%) or trunk broken (43.7%) were common and uprooted (12.2%) was an uncommon type of death of canopy trees. These figures indicate that general features of gap characteristics in this forest type are the low proportion of gap area and the high proportions of small gap size and multiple-tree gap formation. In general, shade-tolerantAbies frequently, andTsuga, infrequently, regenerate in gaps from advance regenerations recruited before gap formation, whilePicea and shade-intolerantBetula possibly regenerate in gaps from new individuals recruited after gap formation. Gap successors of conifers occurred in a wide range of gap size and did not show the clear preference to species specific gap size. In old-growth stands without large-scale disturbance (≥0.1 ha in area) of subalpine coniferous forests of central Japan, major tree species may coexist with their different gap-regeneration behaviors and, probably, different life history traits.     

Canopy and age structures of some old sub-boreal Picea stands in British Columbia

Abstract. 14 old, unlogged, Picea-dominated stands in the moist cool Sub-Boreal Spruce biogeoclimatic subzone of central British Columbia, Canada, were sampled to describe canopy heterogeneity, regeneration patterns and tree population age structures. These stands are composed of Picea engelmannii × glauca hybrids, Abies lasiocarpa and lesser amounts of Pinus contorta and Populus tremuloides, and had survived 124–343 yr since the last stand-destroying wildfire. Canopy cover was patchy and highly variable (ranging from 30.5 % to 86.4 %) but was not significantly related to stand age. Vertical canopy structure was less variable, reflecting the shade-tolerance and live crown ratios (length of live canopy expressed relative to tree height) of component species: 18.8 % for Populus, 20.2 % for Pinus, 46.7 % for Picea and 51.4 % for Abies. Individual stands varied considerably in their population structures and in their stand development trajectories, yet some patterns are evident. Survivors of the initial post-disturbance cohort of trees took 51 to 118 yr (mean = 80, s.d. = 20) to establish. Some stands had all tree species present during stand initiation, while other stands indicated early successional roles for Populus and Pinus, or a late successional role for Abies. Abies recruitment, while often slow in the beginning, occurs uniformly throughout the history of most stands, reflecting the high shade-tolerance of this species. Picea is often recruited in high densities early in stand development, and then (after long periods of exclusion) may be displaced by Abies in some stands but maintains itself in others. Minor, single-tree disturbances (due to bark beetles, root rot, and windthrow) were important in accelerating the reinitiation of Picea in the understory. Results thus suggest that stands from this region can be self-perpetuating in the absence of fire. Yet, post-fire tree populations still clearly dominate these spruce-fir forests, for only the oldest stand had greater basal area in the replacement cohort than in the initial cohort.     

Regeneration in subalpine coniferous forests

The regeneration process of a subalpine coniferous forest, a mixed forest ofTsuga diversifolia (dominant species),Abies veitchii, Abies mariessi, andPicea jezoensis var.hondoensis, was studied on the basis of annual ring data. The age class distribution was discontinuous and four age groups occurred in the study plot (30m×30m). The canopy layer was a mosaic of patches (83.8–133.7 m² patch area), which had different mean ages. The recruitment of canopy trees was carried out only by advance regeneration in the plot. The diameter growth ofAbies andPicea exceeded diameter growth ofTsuga in the gap.Abies lived for 200–300 years and their trunks were susceptible to heart rot.Picea lived for 300–400 years andTsuga for more than 400 years. The regeneration process derived from the analysis of the plot consisted of three phases leading to the development of a even-aged patch; (1) the establishment of saplings before a gap opening, (2) the opening of a gap in the canopy and repair of the canopy by advance regenerated saplings dominated by rapid growth species,Abies andPicea, and (3) the dying off of canopy trees as each species reached the end of its life-span, resulting in pure patches of long-livedTsuga.     

Stand structure and regeneration in a Kamchatka mixed boreal forest

Abstract. A 1‐ha plot was established in a Betula platyphylla‐Picea ajanensis mixed boreal forest in the central Kamchatka peninsula in Russia to investigate stand structure and regeneration. This forest was relatively sparse; total density and stand basal area were 1071/ha and 25.8 m²/ha, respectively, for trees > 2.0 cm in trunk diameter at breast height (DBH). 25% of Betula regenerated by sprouting, and its frequency distribution of DBH had a reverse J‐shaped pattern. In contrast, Picea had a bimodal distribution. The growth rates of both species were high, reaching 20 m in ca. 120 yr. The two species had clumped distributions, especially for saplings. Betula saplings were not distributed in canopy gaps. Small Picea saplings were distributed irrespective of the presence/absence of gaps, while larger saplings aggregated in gaps. At the examined spatial scales (6.25–400 m²) the spatial distribution of Betula saplings was positively correlated with living Betula canopy trees and negatively with dead Picea canopy trees. This suggests that Betula saplings regenerated under the crowns of Betula canopy trees and did not invade the gaps created by Picea canopy trees. The spatial distribution of Picea saplings was negatively correlated with living and dead Betula canopy trees and positively with dead Picea canopy trees. Most small Picea seedlings were distributed under the crowns of Picea trees but not under the crowns of Betula trees or in gaps. This suggests that Picea seedlings establish under the crowns of Picea canopy trees and can grow to large sizes after the death of overhead Picea canopy trees. Evidence of competitive exclusion between the two species was not found. At a 20 m × 20 m scale both skewness and the coefficient of variation of DBH frequency distribution of Picea decreased with an increase in total basal area of Picea while those of Betula were unchanged irrespective of the increase in total basal area of Betula. This indicates that the size structure of Picea is more variable with stand development than that of Betula on a small scale. This study suggests that Betula regenerates continuously by sprouting and Picea regenerates discontinuously after gap formation and that the species do not exclude each other.     

Structure of a pristine Picea abies forest in northeastern Europe

Abstract. The forest structure in a large, relatively homogeneous area of pristine Picea abies taiga in the southern boreal region west of the Ural mountains was studied along four 500-m long transects. P. abies dominated the forest in association with Abies sibirica and Betula spp. The mean volume of living trees was 216 m³/ha. This value varied among the four transects, from 138 - 252 m³/ha. Mean density of trees (DBH > 1 cm) (and variation over the transects) was 2 064/ha (1670 - 2710). Living trees classified as dying or seriously damaged made up 2.9 (2.5 - 3.5) % of the living tree volume. Betula was an important canopy component and made up 16% of the living tree volume. The number of dead standing trees varied from 195 - 325/ha, corresponding to a volume of 10.8 - 70.7 m³/ha. The density of trees with a broken stem was 90 - 170/ha and their estimated volume 7.6 - 41.3 m³/ha. Standing dead trees and trees with broken stems represented 10.4 and 8.9% of the total standing tree volume (living + dead), respectively. The mean volume of decaying logs on the forest floor was 117 (84.4–156.2) m³/ha, corresponding to 54 (35 - 113) % of the living tree volume. The canopy-forming trees were present in the understory as abundant saplings and suppressed individuals. The size distribution of the dominant tree species resembled the inverse J-shape. Generally, the forest was characterized by a high small-scale structural variation and a larger-scale relative homogeneity. This pattern is consistent with forest dynamics where the forest consists of a small-scale mosaic of patches in different stages of recovery following disturbance. Our results suggest that the ecology and dynamics of extensive areas of natural boreal forests can be driven by small-scale disturbance.     

Fire disturbance and forest structure in an old‐growth Pinus ponderosa forest,southern Cascades,USA

Questions: Did fire regimes in old‐growth Pinus ponderosa forest change with Euro‐American settlement compared to the pre‐settlement period? Do tree age structures exhibit a pattern of continuous regeneration or is regeneration episodic and related to fire disturbance or fire‐free periods? Are the forests compositionally stable? Do trees have a clumped spatial pattern and are clumps even‐ or mixed‐age? How might information from this old‐growth forest inform current restoration and management practices? Location: A 235‐ha old‐growth forest in the Ishi Wilderness, southern Cascade Mountains, California. Methods: Age, size, and spatial pattern of trees were quantified in seven stands. Fire history was reconstructed using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing fire history with age, size, and spatial structure of trees and identifying and measuring trees killed by two recent fires. Results: Species composition in plots was similar but density and basal area of tree populations varied. Age structure for P. ponderosa and Quercus kelloggii showed periods of episodic recruitment that varied among plots. Fire disturbance was frequent before 1905, with a median period between fires of 12 years. Fire frequency declined after 1905 but two recent fires (1990, 1994) killed 36% and 41% of mostly smaller diameter P. ponderosa and Q. kelloggii. Clusters of similar age trees occurred at scales of 28‐1018 m² but patches were not even‐aged. Interactions between tree regeneration and fire promoted development of uneven age groups of trees. Conclusions: Fire disturbance strongly influenced density, basal area, and spatial structure of tree populations. Fire exclusion over the last 100 years has caused compositional and structural changes. Two recent fires, however, thinned stands and created gaps favorable for Q. kelloggii and P. ponderosa regeneration. The effects of infrequent 20th century fire indicate that a low fire frequency can restore and sustain structural characteristics resembling those of the pre‐fire suppression period forest.     

Gap characteristics and gap regeneration in a subalpine coniferous forest on Mt Ontake,central Honshu,Japan

Gap characteristics and gap regeneration were studied in three mature stands belonging to different community types in a subalpine coniferous forest on Mt Ontake, central Honshu, Japan. Gap disturbance regimes were remarkably similar among stands studied; percentage gap area to surveyed area, gap density and mean gap size were 7.3–8.5%, 17.8–20.0 ha⁻¹ and 40.8–42.5 m², respectively. The gap size class distributions were also similar and showed a strong positive skewness with a few large and many small gaps; gaps <40m² were most frequent and gaps >200m² were rare. Forty-five to 66% of gaps were due to the death of single canopy trees. Canopy trees more often died leaving standing dead wood (40–5.7%) or broken trunks (43–49%). Shade tolerantAbies mariesii andAbies veitchii, frequently, andTsuga diversifolia, less frequently, regenerated in gaps, from advance regenerations recruited before gap formation.Picea jezoensis var.hondoensis may regenerate in gaps, from new individuals recruited after gap formation. The breakage of denseSasa coverage and the mineral soils exposed by the uprooted plants that form gaps might provide regeneration opportunities for shade intolerantBetula. Of the important species limited to the ridge site,Chamaecyparis obtusa, frequently, andThuja standishii, infrequently, regenerated from plants alreadyin situ. Regeneration ofPinus parviflora was not seen. Based on the gap characteristics and gap regeneration behaviour of each species described, stand dynamics in each stand are discussed.     

Application of a rapid thin section method for observations on decomposing litter in mor humus form in a subalpine coniferous forest

Morphological changes in the decomposing litter ofAbies spp. andBetula spp. in a mor humus form were studied by a rapid thin section method. According to the morphological characteristics, the epidermis, mesophyll and vascular bundleof Abies needle litter were classified into four types: (i) newly fallen; (ii) slightly decomposed; (iii) moderately decomposed; and (iv) greatly decomposed. The distribution of these tissue types along the profile of the forest floor was then investigated. The morphological changes in other litter types, such as branches, scales andBetula leaves during decomposition were observed directly with microscope and electron microscope. Five vertical thin sections and 80 horizontal thin sections were used for these observations and investigations. the decomposition ofAbies litter was slower than that ofBetula litter. The relative decomposition rate of the tissues was in the order of: mesophyll>vascular bundle >epidermis inAbies needles; mesophyll≥epidermis>vascular bundles inBetula leaves; and inner bark >xylem>outer bark in bothAbies andBetula branches. The last remains of the litter were usually stomata, segments of seminiferous scale and outer bark ofAbies. The decomposition of plant litter occurred mainly within the L and F layers of the soil (0–5 cm in depth).Abies needles andBetula leaves completely disappeared at depths of 0–6 cm and 0–4 cm, respectively. Branches disappeared within the top of 5 cm and 6–8 cm forBetula and forAbies, respectively. The scales ofAbies were most slowly decomposed in the soil layers.     

Altitudinal changes in carbon storage of temperate forests on Mt Changbai, Northeast China

A number of studies have investigated regional and continental scale patterns of carbon (C) stocks in forest ecosystems; however, the altitudinal changes in C storage in different components (vegetation, detritus, and soil) of forest ecosystems remain poorly understood. In this study, we measured C stocks of vegetation, detritus, and soil of 22 forest plots along an altitudinal gradient of 700–2,000 m to quantify altitudinal changes in carbon storage of major forest ecosystems (Pinus koraiensis and broadleaf mixed forest, 700–1,100 m; Picea and Abies forest, 1,100–1,800 m; and Betula ermanii forest, 1,800–2,000 m) on Mt Changbai, Northeast China. Total ecosystem C density (carbon stock per hectare) averaged 237 t C ha⁻¹ (ranging from 112 to 338 t C ha⁻¹) across all the forest stands, of which 153 t C ha⁻¹ (52–245 t C ha⁻¹) was stored in vegetation biomass, 14 t C ha⁻¹ (2.2–48 t C ha⁻¹) in forest detritus (including standing dead trees, fallen trees, and floor material), and 70 t C ha⁻¹ (35–113 t C ha⁻¹) in soil organic matter (1-m depth). Among all the forest types, the lowest vegetation and total C density but the highest soil organic carbon (SOC) density occurred in Betula ermanii forest, whereas the highest detritus C density was observed in Picea and Abies forest. The C density of the three ecosystem components showed distinct altitudinal patterns: with increasing altitude, vegetation C density decreased significantly, detritus C density first increased and then decreased, and SOC density exhibited increasing but insignificant trends. The allocation of total ecosystem C to each component exhibited similar but more significant trends along the altitudinal gradient. Our results suggest that carbon storage and partitioning among different components in temperate forests on Mt Changbai vary greatly with forest type and altitude.     

Tree mortality and effects of release from competition in an old‐growth Fagus‐Abies‐Picea stand

Abstract. In a montane mixed Fagus‐Abies‐Picea forest in Babia Gora National Park (southern Poland), the dynamics of an old‐growth stand were studied by combining an 8‐yr annual census of trees in a 1‐ha permanent sample plot with radial increments of Abies and Picea growing in the central part of the plot. The mortality among the canopy trees was relatively high (10% in 8 yr), but the basal area increment of surviving trees slightly exceeded the losses caused by tree death. DBH increment was positively correlated with initial diameter in Abies and Picea, but not in Fagus. For individual trees smaller than the median height, basal area increment was positively related to the basal area of old snags and the basal area of recently deceased trees in their neighbourhood, but negatively related to the basal area of live trees. Dendrochronological analysis of the past growth patterns revealed numerous periods of release and suppression, which were usually not synchronized among the trees within a 0.3 ha plot. The almost normal distribution of canopy tree DBH and the small number of young individuals in the plot indicated that stand dynamics were synchronized over a relatively large area and, hence, were consistent with the developmental phase concept. On the other hand, the lack of synchronization among periods of growth acceleration in individual mature Abies and Picea trees conforms more closely to the gap‐dynamics paradigm.     

Tree replacement patterns in subalpine Abies-Betula forests,Wolong Natural Reserve,China

Vegetation in canopy gaps of two old-growth Abies-Betula forest stands, one with bamboo the other without, was measured. The structure of gap vegetation at each site was used to derive tree replacement probabilities. Transition probabilities indicate different tree replacement trends in forests with bamboo compared to those without. Projected compositions show Betula to be the most abundant species in bamboo stands while Abies remains most abundant where bamboo was absent. A dense bamboo sward seems to reduce the probability of Abies filling gaps by inhibiting establishment and growth of seedlings. Bamboo preempts space after canopy gap formation by increasing shoot production which reduces opportunities for establishment and growth of other woody species. Differences in dispersal ability and longevity of Abies and Betula appear to be important factors contributing to their coexistence forests with a small canopy gap disturbance regime.     

Five centuries of changing forest vegetation in the Northeastern United States

Maps of physiognomic classes of vegetation in the northeastern UnitedStates before European human impact and for the late 20th century, based onpollen data, closely resemble each other, indicating a robust pattern in theface of the novel and extensive disturbances of the last 500 years. On the otherhand, species abundances have changed considerably, with today's vegetationhaving less Fagus L. and TsugaCarrière and more Betula L.,Picea A. Dietr., and Abies Miller.Picea and Castanea Miller increasedfrom 1700 to 1900, but have decreased in this century. The difference betweenthe main physiognomic classes has remained strong, however, with continuouslymore Picea, Abies andBetula in the north and more QuercusL. and Carya Nutt. in the south, indicating the dominanceof climate in regulating the relative abundances of these genera. Thevegetational patterning within these broad classes has changed in some placesand remained stable in others, suggesting differences in local factors thatdetermine relative species abundances. The influence of slight altitudinalgradients, for example, may influence forest composition only after severalgenerations of trees, while substrate differences may be apparent in the firstforest that regenerates after agricultural abandonment or logging. Forests oftoday, therefore, reflect complex interactions of disturbance and environment,and may be in equilibrium with some but not all features of their currentenvironments.     

Successional trends 219 years after fire in an old Pinus sylvestris stand in northern Sweden

Abstract. We present results from repeated analyses (1962, 1993) of a permanent plot established in 1947, combined with retrospective stand age structure data, in an old Pinus sylvestris stand in Muddus National Park, northern Sweden. The study points towards a successional pathway governed by concurrent disturbance effects of climate variability, reindeer grazing and fire. This is intermediate to the two often advocated ideas on dynamics in boreal forests, that is, one of disturbance-related tree regeneration/mortality and one of continuous regeneration. When the plot was established in 1947 the tree layer (> 1.3 m) consisted of 300 individuals/ha of P. sylvestris and 62/ha of Betula pubescens. Subsequently the stand has become more dense and the species dominance has shifted. In 1993, 362 P. sylvestris and 62 Picea abies individuals were present per ha, while no Betula individuals were found. The number of dead trees increased from zero in 1947 to 200/ha (Pinus) in 1993. Pinus was also the most common species in the sapling layer (< 1.3 m) throughout the study period, though the number dropped from 8912/ha in 1947 to 51% in 1993. Dead saplings decreased from 2650/ha in 1947 to ca. 50% in 1962, and only 9% in 1993. Temporal variations in mortality and sapling mean height coincided with variations in snow depth, indicating a critical period in sapling development when saplings are exposed at the snow/atmosphere interface. The number of living Picea saplings increased slowly until 1993; no dead saplings were found. Most Pinus recruited shortly after the 1774-fire, and during the second half of the 1900s. The major part of the spruce regeneration took place during the later half of the 1900s. No successful Betula recruitment has occurred after the 1930s, and no live Betula were present in 1993, which might be explained as an effect of increased reindeer browsing – the reindeer stock has grown by 50% since 1961. Although subjected to high mortality, Pinus regenerated and maintained a seedling/sapling bank. In this way Pinus remained dominant in the tree layer after more than 200 post-fire years. The importance of the shade-tolerant Picea has slowly increased, while Betula has died off. Thus, even after 219 yr since fire there is an early successional trend in the stand. This suggests that an increased chronic disturbance (grazing/browsing by reindeer) has partly succeeded earlier discrete fire-disturbance events, and maintained a continuous seedbed favouring the shade-intolerant pine recruitment.     

Regeneration in subalpine coniferous forests

Death of canopy trees when gaps are formed was studied in a subalpine coniferous forest, central Japan, which was composed ofAbies, Tsuga, Picea, Betula, andSorbus. Typhoons were considered to be the most important cause of the death of canopy trees. The degree of disturbance in each of 16 plots (20 m×20 m) was represented by the percentage of the total basal area of dead trees to that of living and dead canopy trees (disturbance magnitude; MAG). The mortality of canopy trees increased as their dbh increase in the plots of lower MAG than 90%. The mortality varied among genera, andTsuga was characterized as having lower mortality than that of the other conifers. 418 dead trees were observed. The standing dead trees made up 10.7% of the trees, the stem broken trees 46.7%, and the uprooted trees 42.2%. The stem breaking was most frequent inAbies, and the uprooting was most frequent inTsuga, Picea, andBetula. Undeveloped forests, which have the L-shaped dbh distribution, were destroyed only in high degree (70%<MAG), while developed forests were destroyed in various degrees (30%<MAG<100%). The percentage of uprooted trees in basal area decreased with the development of the forest, from 60% to 10%.     

A new Late-glacial and Holocene record of vegetation and fire history from Lago del Greppo, northern Apennines, Italy

Detailed Late-glacial and Holocene palaeoenvironmental records from the northern Apennines with a robust chronology are still rare, though the region has been regarded as a main area of potential refugia of important trees such as Picea abies and Abies alba. We present a new high-resolution pollen and stomata record from Lago del Greppo (1,442 m a.s.l., Pistoia, northern Apennines) that has been dated relying on 12 terrestrial plant macrofossils. Late-glacial woodlands became established before 13000 cal b.p. and were dominated by Pinus and Betula, although more thermophilous taxa such as Quercus, Tilia and Ulmus were already present in the Greppo area, probably at lower altitudes. Abies and Picea expanded locally at the onset of the Holocene at ca. 11500 cal b.p. Fagus sylvatica was the last important tree to expand at ca. 6500 cal b.p., following the decline of Abies. Human impact was generally low throughout the Holocene, and the local woods remained rather closed until the most recent time, ca. a.d. 1700–1800. The vegetational history of Lago del Greppo appears consistent with that of previous investigations in the study region. Late-glacial and Holocene vegetation dynamics in the northern Apennines are very similar to those in the Insubrian southern Alps bordering Switzerland and Italy, across the Po Plain. Similarities between the two areas include the Late-glacial presence of Abies alba, its strong dominance during the Holocene across different vegetation belts from the lowlands to high elevations, as well as its final fire and human-triggered reduction during the mid Holocene. Our new data suggest that isolated and minor Picea abies populations survived the Late-glacial in the foothills of the northern Apennines and that at the onset of the Holocene they moved upwards, reaching the site of Lago del Greppo. Today stands of Picea abies occur only in two small areas in the highest part of the northern Apennines, and they have become extinct elsewhere. Given the forecast global warming, these relict Picea abies stands of the northern Apennines, which have a history of at least 13,000 years, appear severely endangered.     

The relationship between fire history and an exotic fungal disease in a deciduous forest

Exotic diseases have fundamentally altered the structure and function of forest ecosystems. Controlling exotic diseases across large expanses of forest has proven difficult, but fire may reduce the levels of diseases that are sensitive to environmental conditions. We examined Cornus florida populations in burned and unburned Quercus–Carya stands to determine if burning prior to anthracnose infection has reduced the impacts of an exotic fungal disease, dogwood anthracnose, caused by Discula destructiva. We hypothesized that fire has altered stand structure and created open conditions less conducive to dogwood anthracnose. We compared C. florida density, C. florida health, and species composition and density among four sampling categories: unburned stands, and stands that had burned once, twice, and 3 times over a 20-year period (late 1960s to late 1980s). Double burn stands contained the greatest density of C. florida stems (770 stems ha⁻¹) followed by triple burn stands (233 stems ha⁻¹), single burn stands (225 stems ha⁻¹) and unburned stands (70 stems ha⁻¹; P < 0.01). We observed less crown dieback in small C. florida trees (<5 cm diameter at breast height) in burned stands than in unburned stands (P < 0.05). Indicator species analysis showed that burning favored species historically associated with Quercus–Carya forests and excluded species associated with secondary succession following nearly a century of fire suppression. Our results suggest that fire may mitigate the decline of C. florida populations under attack by an exotic pathogen by altering forest structure and composition. Further, our results suggest that the burns we sampled have had an overall restorative effect on forest communities and were within the fire return interval of the historic fire regime. Consequently, prescribed fire may offer a management tool to reduce the impacts of fungal disease in forest ecosystems that developed under historic burning regimes.     

Revegetation process on a subalpine mudflow

The process of revegetation on a subalpine mudflow was analyzed on the basis of the vegetation-soil relationship of the revegetated stands, and of the age structure and height growth process of the trees. Based on evidence derived from five independent sources, it was concluded that this mudflow was caused by a typhoon in 1959. The revegetated stands were classified into three vegetation types. AnAlnus maximowiczii stand was established where the surface soil was unstable; aBetula ermanii stand was established where the degree of disturbance by the mudflow was small; anAlnus-Betula stand was the intermediate type and occupied most of the area on the mudflow.Alnus had colonized concentrically 4–6 yr after the mudflow, andBetula and conifers had followed 1–2 yr later. As revegetation proceeded, the trees became differentiated into two populations, a canopy population and a suppressed one. The former was dominated mainly byAlnus and the latter byBetula and conifers. This differentiation was caused by the faster growth ofAlnus during the initial years of colonization. SomeBetula trees have recently grown faster thanAlnus, suggesting future replacement in dominance. Conifers have grown slowly even during recent years; the highly oligotrophic condition of the soil appears to have been responsible for this delayed growth.     

Stand dynamics of Pinus flexilis-dominated subalpine forests in the Colorado Front Range

Population age structure and succession were investigated in subalpine forests in the Colorado Front Range dominated by Pinus flexilis (limber pine). Age, size, and spatial data were collected from three recent burns (<100 yr old), six ca. 240 year-old post-fire stands, and two old-growth stands (individuals > 400 yr old). The sequence of colonization of now extant trees on these post-fire sites appeared to be consistent: first Pinus flexilis, then Picea engelmannii (Engelmann spruce), and later Abies lasiocarpa (subalpine fir) with a delay between the first Pinus flexilis and Abies of as long as 140 yr. The advantage of Pinus flexilis on post-burn sites can be attributed to avian seed dispersal and the exceptional drought tolerance of its seedlings. The three recent burns were not extensive, and the delay in establishment of Picea and Abies appeared to be limited by harsh site conditions rather than lack of seed dispersal. Spatial analysis indicated a consistent, although sometimes weak, attraction between Pinus flexilis and Picea and Pinus flexilis and Abies at a scale of 1–4 m, suggesting that Pinus flexilis may facilitate establishment of Picea and Abies seedlings by providing shade or protection from wind. On xeric to slightly xeric sites, Pinus flexilis appeared to form broadly even-aged, non-regenerating populations that were gradually being replaced by Picea and Abies. Replacement is proceeding at a faster rate on the least xeric sites (north aspects, valley bottoms) compared to the most xeric sites (south aspects). On the most extreme sites, Pinus flexilis formed all-aged, self-maintaining populations with no evidence of replacement by Picea and Abies. In these old-growth forests with occasional trees aged at > 1300 yr, recruitment is continuous or episodic.     

Present and past old‐growth forests of the Lake Tahoe Basin,Sierra Nevada,US

Abstract. We described 38 relictual old‐growth stands – with data on the mortality, regeneration, floristic richness, fuel load and disease incidence in our study area in the Tahoe Basin of California and Nevada. The stands are within the lower and upper montane zones (1900–2400 m a.s.l.) and they are rare, occupying < 2% of the land in the Basin's watershed. Correlation matrices and ANOVAs of forest types and conifer species with environmental gradients revealed significant relationships with elevation, distance east of the Sierran crest, slope aspect, annual precipitation, date of complete snow melt, litter depth and degree of soil profile development. Pathogens, parasites and wood‐boring insects were present on 23% of living trees; 16% of all trees were dead. We compared these stands to a reconstruction of pre‐contact Basin forests and to ecologically analogous old‐growth forests of Baja California that have never experienced fire suppression management. Currently, overstorey trees (> 180 yr old) in the Basin stands have ca. 33% cover, 54 m².ha^‐1 basal area and 107 individuals.ha^‐1, values very similar to reconstructions of pre‐contact Basin forests and to modern Baja California forests. Understorey trees (60–180 yr old), however, are several times more dense than historic levels and species composition is strongly dominated by A. concolor, regardless of the overstorey composition. The ratio of Pinus: Abies has increased – and the age structure of extant stands predicts that it will continue to increase – from approximately 1:1 in pre‐contact time to 1:7 within the next century. Disease incidence and mortality in Baja forests were lower. Although we quantitatively defined current Basin old‐growth forests – in terms of stand structure – we realize that our definition will differ from that of both past and future old‐growth forests unless management protocols are changed.     

Plastic Response of Crown Architecture to Crowding in Understorey Trees of Two Co-dominating Conifers

Crown architecture and growth rate of trunk height, trunk diameterand lateral branches of understorey trees (5–10m tall)were compared between two co-dominating conifers,Abies sachalinensisandPiceaglehnii, in relation to the index of local crowding intensity,W,represented as a function of density, distance and basal areaof taller neighbours. For the two conifers, the growth of trunkheight and diameter was decreased and crowns became flat withincreasingW, keeping crown projection area. Self-pruning oflower branches was more intense inAbiesthan inPiceain crowdedconditions, while both conifers showed similar crown forms inless crowded conditions. These results suggest that the growthin lateral branches exceeded that in height in crowded conditions,especially inAbies. Tree age of both conifers increased withincreasingW, resulting from the low growth rate in crowded conditions.The age of the longest and lowest branch ofPicea, up to 150years, was positively correlated with the tree age ranging from70 to 250 years, whereas that ofAbieswas constant at around30 years irrespective of tree age varying from 40 to 140 years.This result agrees with the observation that agedAbieshad moreflat-shaped crowns than in agedPiceain crowded conditions. Theseresults suggest that each conifer adapted to crowding in differentways: high elongation of branches with high turnover rate forAbiesandviceversaforPicea. Abies; crown form; neighbourhood interference; Picea; plasticity