Effect of bryophytes on survival of conifer seedlings in subalpine forests of central Japan

Bryophytes form the major seedbed for coniferous trees in the subalpine forests of central Japan. Field experiments were conducted on the floor of a closed stand dominated byTsuga on Mt Fuji to examine the seedling survival ofAbies veitchii andTsuga diversifolia in seven substrate types in relation to the morphological characteristics of the seeds and seedlings. NeitherAbies norTsuga seedlings could survive on dwarf-bamboo litter beyond 2 years after the seed rain.Abies seedlings survived not only in all types of bryophyte communities but also in larch litter. In contrast, the survival ofTsuga seedlings was restricted to communities of smaller-statured bryophytes found on logs. The seeds and 1st year seedlings ofA. veitchii were larger than those ofT. diversifolia. Abies seedlings produced hypocotyls taller than any bryophyte community and radicles which were long enough to penetrate into the moist humus layer, whereasTsuga seedling radicles presumably penetrate into humus only in bryophyte communities forming a thin mat on logs. The effect of bryophytes on survival of tree seedlings therefore depends on the relationships between the morphology of seeds and seedlings and the structure of bryophyte communities.     

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 dynamics of wave-typeAbies sachalinensis coastal forest

Stand structure and regeneration pattern were examined inAbies sachalinensis coastal forest in northern Hokkaido. In the forest a similar phenomenon to the wave regeneration in subalpine forests has been found. Wave regeneration has been reported for montaneAbies forests in central Japan and North America. Differences and similarities between wave-type stands in this coastal forest and wave-regenerated montane forests were clarified. The shift of dead tree zone, stand structure and regeneration pattern in wave-type stands are the same as in subalpine wave-regenerated forests. High density of individuals is considered to be an internal factor which causes stand-level dieback and also enables the stands to persist in the severe environment in both forests. A difference between wave-regenerated forests andA. sachalinensis wave-type stands is the number of dead tree zones, which is only one in wave-type stands. Changes of regeneration patterns ofAbies sachalinensis with environmental gradient from seaward to inland were related to this difference.     

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.     

Vertical vegetation zones along 30° N latitude in humid East Asia

Structural changes in altitudinal vegetation zones along a 30° N parallel were studied based on vegetation data from 20 mountains in East Asia, from 85° E to 130° E longitude. The altitude of comparable vegetation zones showed a sharp increase of 1400–1900 m from east to west. Forest limit reached an altitude of 4400–4600 m in the eastern Tibetan Plateau, being the highest forest limit in the world. The limidng factor for the upper limit of a vegetation zone was different in the east and west. Low temperature in winter controlled the upward distribution of the evergreen broadleaf forest in the east, whereas the limiting factor was growing season warmth in the west. A close correlation was found between the climatic indices and annual range of monthly mean temperature (ART) at the upper limit of a vegetation zone.Component genera of each vegetation zone along the 30° N parallel were analyzed, and it was found that the alternation of component genera from east to west was much more apparent in cool-temperate forests, reflecting their response to macrotopography and air masses. The distribution of Fagus extended into winter-cold regions, whilst Tsuga occurred principally in oceanic and warm climates. The northern limit of Tsuga corresponded well to an ART isotherm of 23 °C and its southern limit coincided with that of Fagus. According to the distribution of Fagus and Tsuga, the cool-temperate forests in East Asia along the 30° N belt were divided into three types: deciduous broadleaf forest (represented by Fagus), mixed forest (dominated by Fagus, Tsuga and others), and mixed evergreen forest (consisting mainly of Tsuga and sclerophyll oaks).     

Succession in sub‐boreal forests of West‐Central British Columbia

Abstract. Structural and compositional changes were analysed over the course of 400+ yr of post‐fire succession in the sub‐boreal forests of west‐central British Columbia. Using a chronosequence of 57 stands ranging from 11 to 438 yr in age, we examined changes in forest structure and composition with complementary PCA and DCA ordination techniques. To determine stand ages and timing of tree recruitment, approximately 1800 trees were aged. Most early successional forests were dominated by Pinus contorta, which established rapidly following fire. Abies lasiocarpa and Picea glauca × engel‐mannii were also able to establish quickly, but continued to establish throughout the sere. Few Pinus contorta survived beyond 200 yr, resulting in major changes in forest structure. In some stands P. contorta never established, which led to considerable variation among stands less than 200 yr old. The oldest forests converged on dominance by Abies lasiocarpa. Vascular plant diversity decreased during succession whereas canopy structure became more complex as gap dynamics developed. Although these sub‐boreal forests contain few tree species, successional changes were pronounced, with structure changing more than composition across the chronosequence.     

The dynamics of old-growth Pseudotsuga forests in the western Cascade Range,Oregon, USA

Size and age structure, spatial analysis, and disturbance history were used to analyse the population structures and regeneration patterns of 8 conifer stands in the central western Cascade Range, Oregon, USA. Variation in forest structure reflected the effects of frequent (20–50 yr) low-intensity fires and treefalls, infrequent (100–200 yr) localised, intense fires, and extensive fires that resulted in stand replacement (every ca 400 yr?).The amount of canopy removed and the size of openings formed by fires and treefalls were important determinants of subsequent forest establishment. Single or several species stands of Pseudotsuga and/or Abies procera, or mixed species stands of Pseudotsuga, Abies procera, Tsuga heterophylla and Abies amabilis established in openings where intense fires had removed most of the canopy trees over several ha. Multi-tiered and multi-aged stands, often containing 400–500 yr-old Pseudotsuga and variously-sized more or less even-aged patches of younger shade tolerant Tsuga heterophylla and/or Abies amabilis, occurred where lower-intensity fires did not kill all overstorey trees or where treefalls occurred after the initial fire.Current regeneration processes are influenced by overstorey composition, the availability and size of canopy openings, and the availability of substrates suitable for regeneration. Tsuga heterophylla and Abies amabilis established under Pseudotsuga menziesii and Abies procera canopies and in small canopy openings (<400 m²) created by windfalls, but rarely under Tsuga. Down logs and stumps were favoured establishment sites for Tsuga.The disturbance regime of fires of low-to moderate-intensity, windfalls, and occasional fires that result in extensive stand replacement contrasts with the pattern of infrequent, catastrophic disturbances proposed for other areas of the Pacific Northwest. Although fires at stand establishment commonly determine much of the composition, structure, and subsequent stand development, canopy replacement by shade tolerant species occurs as the different life histories of the species are expressed in response to various disturbances differing in intensity and frequency. Such a non-equilibrium view of vegetation change is consistent with many other fire-dominated forests of the western United States.     

Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan

To evaluate the effects of canopy gaps and forest floor microsites (soil, fallen logs, root-mounds, buttresses and stumps) on regeneration of subalpine forests, the gap regeneration and seedling occurrence of conifers (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis and Tsuga diversifolia) were studied in two stands of a subalpine old-growth forest, central Japan. The percentage of gap area to total surveyed area was 11.2–11.3% in the stands. Gap regeneration was not common for P. jezoensis var. hondoensis and T. diversifolia. In contrast, gap regeneration by advanced regeneration was relatively common for Abies. Seedling occurrence of P. jezoensis var. hondoensis and T. diversifolia was restricted on elevated surfaces such as stumps and root-mounds, while Abies seedlings could occur on soil as well as on elevated surfaces. Rotten stumps were the most favorable microsites for conifer seedling occurrence, which covered small area in the forest floor. Although canopy gaps were not always favorable for seedling occurrence, all conifer seedlings were larger under canopy gaps than under closed canopy. Canopy gaps and forest floor microsites clearly affected seedling occurrence and growth of conifers. This suggests that regeneration of conifers is related to the difference of growth advantage under canopy gaps and favorable microsites for seedling occurrence.     

The population biology of Bromus tectorum in forests: distinguishing the opportunity for dispersal from environmental restriction

Summary With increasing elevation and corresponding changes in the macroclimate, forest zones in the Intermountain Region of western North America are often dominated in turn by Pinus ponderosa, Pseudotsuga menziesii, Abies grandis, an Thuja plicata. Bromus tectorum (cheatgrass), and introduced annual grass now abundant in the Region's steppe, is uncommon in mature stands representative of these forest zones. In order to determine whether B. tectorum is largely excluded from these forests by insufficient seed dispersal or environmental restriction(s), the grass's demography was compared in each of four years among populations experimentally-introduced into mature forests. The number of recruits did not differ among the Pinus, Pseudotsuga, and Abies sites; recruitment was however significantly lower on the coolest site dominated by Thuja. Emergence in both the low elevation Pinus and Pseudotsuga sites was about the same in autumn, winter, and spring. In the cooler, moister Abies and Thuja sites, emergence was limited to autumn and early winter. Survival in these forest sites ranged between 0 and 87%. The percentage of the total population to survive until harvest was highest in the Pseudotsuga site, intermediate in the Pinus and Abies sites, and lowest in the Thuja site. Compared with B. tectorum in the steppe, the surviving plants were small, and few produced seeds. All parents were members of either the autumn or winter cohort, and most parents produced only one seed. No seeds were produced at the Thuja site. Although phenotypic plasticity apparently contributes to the wide ecological amplitude of this grass, its growing season on these sites in most years is too short for it to reproduce. Consequently, these forest zones broadly define the current environmental limits to the distribution of cheatgrass in this portion of its new geographic range.     

Conservation value of timber quality versus associated non-timber quality stands for understory diversity in Nothofagus forests

Conservation strategies of forested landscapes must consider biodiversity of the included site types, i.e. timber-quality forests and associated non-timber-quality stands. The objectives were to characterize forest overstory structure in timber-quality versus associated non-timber-quality stands; and to compare their understory communities. Six forest types were sampled in Nothofagus forests of Tierra del Fuego (Argentina): two timber-quality N. pumilio forests, and four associated non-timber-quality stands (edge, N. antarctica, wetlands and streamside forests). Overstory structure and understory vegetation (species richness, frequencies, cover and biomass) were characterized during spring and summer seasons. Analysis of variance and multivariates were carried out. Overstory structure differed across the site types, with higher tree size, canopy closure and tree volume in timber-quality stands. Fifty-one understory plant species were observed, but understory variables varied with site types, especially wetlands (highest native and exotic richness, cover and biomass, and 25% of exclusive species). Site types were grouped in three: N. antarctica stands, streamside stands and the other N. pumilio forests according to multivariate analysis. Forty three percent of plants were distributed in all site types, and all timber-quality forest understory species were present in some associated non-timber-quality stands. Timber-quality N. pumilio forests have a marginal value for understory conservation compared to associated non-timber-quality stands, because these last include all the plants observed in timber-quality forests and also possess many exclusive species. Therefore, protection of associated non-timber-quality stands during forest management planning could increase understory conservation at landscape level, and these could be better reserves of understory diversity than retentions of timber-quality stands.     

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.     

State of fallen logs and the occurrence of conifer seedlings and saplings in boreal and subalpine old-growth forests in Japan

Fallen logs on the forest floor play an important role in tree seedling establishment in boreal and subalpine old-growth forests. We examined the abundance and state (degree of decay, moss coverage on their surfaces) of fallen logs on the forest floor and compared the occurrence of conifer seedlings and saplings on soil and fallen logs in old-growth evergreen boreal (Taisetsu) and subalpine (Yatsugatake) coniferous forest stands in Japan. The forest floors of both stands were covered mostly by soil, and fallen logs covered only a small proportion (4.5% in Taisetsu and 2.5% in Yatsugatake) of the forest floor. In both stands, no seedlings were found on freshly fallen logs. Both moss coverage and the occurrence of seedlings or saplings on fallen logs increased with decay in both stands. In Taisetsu, the occurrence of all conifer seedlings and saplings was, in general, restricted to fallen logs. In contrast, in Yatsugatake, Picea and Tsuga seedlings occurred much more frequently on fallen logs than on soil, whereas Abies seedlings and saplings became established on both soil and fallen logs. Seedlings were taller on decayed fallen logs, but taller saplings were rare on fallen logs in Yatsugatake. Results indicate that species characteristics, the condition of the fallen logs and environmental factors determine the preponderance of seedling and sapling establishment on fallen logs. The condition of fallen logs varies with the degree of decay, and fallen logs that function as seedling establishment sites on the forest floor are thought to be limited by time and space.     

Dynamics in Central European near‐natural Abies‐Fagus forests: Does the mosaic‐cycle approach provide an appropriate model?

Question: The mosaic‐cycle concept of forest dynamics dominates in Central Europe. According to this concept intermediate‐scale disturbances only accelerate the forest break‐up under existing cycles of forest development. Is such an approach correct, or should new developmental cycles be elaborated for intermediate‐scale disturbances? Location: Near‐natural Abies alba ‐ Fagus sylvatica forests in the ?wiétokrzyski National Park in Central Poland. In these forests intermediate‐scale disturbances occurred between 1970 and 1990. Methods: Data were collected twice in areas surrounding 212 permanent sample points (in 1994 and 2004). Two increment cores were taken from 259 sample Abies trees. The effect of intermediate‐scale disturbances on radial increment of Abies was assessed. Probabilities of stand transition during a 10‐year period between individual stages and phases of development of the mixed forest were calculated. The development stages and phases were arranged into hypothetical succession series of successive changes. Results: In 1994 70 stands and in 2004, 47 stands representing stages and phases containing the older generation formed by trees > 100–150 years were found. Also, in 1994 142 and in 2004, 165 stands representing stages and phases containing the younger generations only, formed by trees < 100‐150 years, were recorded. Stages and phases containing only younger generations do not occur in the existing forest development cycle which does not consider the influence of intermediate‐scale disturbances separately. Two developmental cycles, which take into account the presence of the older generation and the younger generations only (under conditions of the occurrence of intermediate‐scale disturbances), are proposed. Conclusion: The mosaic‐cycle concept of forest dynamics can be used to analyse the dynamics of Central European near‐natural mixed‐species forests, but new developmental cycles should be elaborated for intermediate‐scale disturbances.     

Plant diversity in managed sal (Shorea robusta Gaertn.) forests of Gorakhpur, India: species composition, regeneration and conservation

The sal (Shorea robusta) forest vegetation of Sohagibarawa Wildlife Sanctuary, Gorakhpur, India was analysed to assess plant diversity, regeneration pattern and the status of species conservation. A total of 208 plant species representing 165 genera and 72 families were recorded. Species richness, mean density and basal area of individuals in the observed forest were compared with those of other sal-dominated forests of India. The sal forest was rich in Papilionaceae (23 species), which contributed maximally to the total number of individuals of <30 cm girth. After sal, density was maximal for a leguminous shrub, Moghania chappar. In addition to the usual recruitment by seed, a number of species also showed non-seed regeneration through storage roots, sprouts or ramet proliferation. The individuals regenerating as sprouts from underground stem or storage organs contributed significantly to the sum total of individuals/ha. As much as 45.5% of the total individuals were of ramet origin and shared 10.6% of the total species richness of the forest. In stands facing moderate to low disturbance, thickets of dense entangled mass of vegetation, predominantly composed of thorny lianas, were identified that usually contained less common and rare species like Rauwolfia serpentina, Desmodium latifolium, Crotalaria alata and Gloriosa superba in addition to the frequent ones. These thickets help to conserve the special habitat conditions and provide protection for natural regeneration of several infrequent and rare plant species and thus contribute towards the maintenance and in situ conservation of overall diversity of recurrently disturbed forest vegetation.     

Picea-Abies forests in the highlands of Northern Alberta

Summary Thirty stands of Picea-Abies dominated forest in highland areas of northern Alberta were studied quantitatively. The highlands are cooler and receive more precipitation than the adjacent lowlands in summer. Orthic Grey Luvisols are the most common soil type.Based on morphological criteria the Picea was basically P. glauca, the Abies mostly A. balsamea, and the Pinus nearly all P. contorta var. latifolia. The boreal floristic element in stands increases from west to east and from high to low altitude.Four Picea-Abies community types were recognized using ordination and cluster analysis: Rubus pedatus/Ptilium crista-castrensis, Cornus canadensis-Linnaea borealis, Viburnum edule/Hylocomium splendens, Calamagrostis canadensis. The 4 community types are well correlated with altitude, soil nutrient status, and 2 groups of correlated floristic and structural variables.The floristic and structural affinities of the 4 community types to those of adjacent regions are discussed.The Picea-Abies stands studied are considered to belong to a group of climax community types, with Abies dominance indicated at higher altitudes, shifting to Picea dominance at lower altitudes.Nomenclature follows Moss (1959) for vascular plants, with the exception of Dryopteris (Britton 1972) and Betula (Dugle 1966); Lawton (1971) for mosses; Schofield (1968) for hepatics; and Hale & Culberson (1970) for lichens.We thank E. M. Achuff and T. P. Achuff for field assistance; D. M. Britton, M. Dumais, M. Ostafichuk, W. Peterson and D. H. Vitt for taxonomic assistance; the Dept of Soil Science, University of Alberta and the Research Branch, Agriculture Canada for the use of laboratory equipment, Financial support was received from the Boreal Institute for Northern Studies and the National Research Council of Canada (A-2570 La Roi).     

Are sub-alpine firs evolving towards semelparity?

Summary In sub-alpine forests of Northeast America and Japan pure stands of trees in the genusAbies exhibit wave regeneration. Opportunities for recruitment in such forests are confined to a window in time and space that coincides with the death of an even-aged cohort of adult trees. I suggest that the coincidence of this recruitment window with the death of adults at a predictable age should select for convergence between age at first reproduction and age at death. Ultimately this would lead to the evolution of semelparity. The available evidence supports this hypothesis for wave-regeneratedAbies populations in Japan. A field test of the hypothesis is also suggested.     

Regeneration in fringe mangrove forests damaged by Hurricane Andrew

Mangrove forests along many tropical coastlines are frequently andseverely damaged by hurricanes. The ability of mangrove forests to regeneratefollowing hurricanes has been noted, but changes that occur in vegetationfollowing disturbance by hurricane winds and storm tides have not been studied.We measured changes in plant community structure and environmental variables intwo fringe mangrove forests in south Florida, USA that experienced high windvelocities and storm tides associated with Hurricane Andrew (August1992). Loss of the forest canopy stimulated regeneration via seedlinggrowth and recruitment, as well as resprouting of some trees that survived thehurricane. Initial regeneration differed among species in both forests:Rhizophora mangle L. regenerated primarily via growth ofseedlings present at the time of the hurricane (i.e., release of advancerecruits), but many trees of Avicennia germinans(L.) Stearn and Laguncularia racemosa Gaertn.f.resprouted profusely from dormant epicormic buds. In one forest, which wasformerly dominated by Laguncularia, high densities ofRhizophora seedlings survived the hurricane and grew toform dense stands of saplings and small trees ofRhizophora. In the other forest, there were lowerdensitiesof surviving Rhizophora seedlings (possibly due tohigher storm tide), and extensive bare areas that were colonized byAvicennia, Laguncularia, andherbaceous species. This forest, predominantly Rhizophoraat the time of the hurricane, now contains stands of saplings and small treesofall three species, interspersed with patches dominated by herbaceous plants.These findings indicate that moderately damaged fringe forests may regenerateprimarily via release of Rhizophora advance recruits,leading to single-species stands. In severely damaged forests, seedlingrecruitment may be more important and lead to mixed-species stands.Regeneration of mangrove forests following hurricanes can involve differentpathways produced by complex interactions between resprouting capability,seedling survival, post-hurricane seedling recruitment, and colonizationby herbaceous vegetation. These differences in relative importance ofregeneration pathways, which may result in post-hurricane forestsdifferent from their pre-hurricane structure, suggest that models forregeneration of mangrove forests will be more complex than directregeneration models proposed for other tropical forests whereregeneration after hurricanes is dominated by resprouting.     

Regeneration and coexistence of two Abies species dominating subalpine forests in central Japan

Summary The mechanism of coexistence of the dominant firs Abies veitchii and A. mariesii is described in relation to regeneration patterns for climax subalpine forests of the northern Yatsugatake Mountains, central Honshu, Japan. Two mature stand types, pure conifer stands of Abies spp., and mixed stands of Abies spp. and hardwoods (mainly the birch Betula ermanii), are distinguished. Pure stands are likely to show simultaneous decay, followed by evenaged regeneration of stand-floor seedlings (<20 cm tall), Rapidly growing A. veitchii dominates over A. mariesii in this type of regeneration, which is occasionally invaded by light-demanding Betula. In constrast, mixed stands degenerate rather slowly, followed by the regeneration of Abies from the bank of suppressed saplings (>20 cm tall), which persist only in mixed stands. The more shade-tolerant A. mariesii is supeior in this type of regeneration, while Betula does not succeed, and mixed stands change to pure stands with time. The fact that two patterns of Abies regeneration occur in a certain ratio in the forest is what enables the two Abies species to coexist. A simple dynamical system model supports this conclusion.     

Structural heterogeneity and tree spatial patterns in an old-growth deciduous lowland forest in Cantabria, northern Spain

Old-growth deciduous forests in western Europe, for the most part, consist of small tracts that often may be atypical due to human disturbance, poor soil productivity or inaccessibility. In addition, very little information on tree age distributions, structural heterogeneity and tree spatial patterns appears to be available for west-European forests. Characterization of the structural features of tree populations in these old-growth stands can provide the basis to design conservation plans and also inform on how present forests might look in the absence of human interference. Four old-growth stands in a deciduous forest in the Cantabrian lowlands, northern Spain, were surveyed to determine forest structure and spatial patterns. Live and dead trees were identified, measured and mapped, and live trees were cored for age estimation. Structural heterogeneity was analyzed by means of the spatial autocorrelation of tree diameter, height and age, and the uni- and bivariate spatial patterns of trees were analyzed. The dominant species, Fagus sylvatica and Quercus robur, showed reverse-J shaped size distributions but discontinuous age distributions, with maximum ages of 255–270 yr. Tree ages suggested that the forest was largely modified by past changes in forest-use, especially by temporal variation in grazing intensity. Spatial autocorrelation revealed that former parkland stands were heterogeneous with respect to tree height only, while high forest stands were composed of patches of even-aged and even-sized trees. Young trees were clumped at varying distances and establishment occurred preferentially in canopy gaps, except for Ilex aquifolium that mainly occurred beneath mature Quercus trees. Surviving trees became less intensely clumped in the dominant species, and more strongly clumped in understorey ones, which may have been due to the effects of intraspecific competition and of canopy trees on tree survival, respectively. The spatial associations between species varied within the forest, probably as a consequence of specific establishment preferences and competitive interactions.     

Age structure and dynamics of Patagonian beech forests in Torres del Paine National Park,Chile

This study documents the stem size and age-structure in forests dominated by different species of Nothofagus in Torres del Paine National Park (51° S), in the Chilean Patagonian region. We also explored the relationship between the various types of Nothofagus forest and postglacial succession. Pioneer stands on moraine fields 1–2 km of the glacier front are dominated by Nothofagus betuloides and Nothofagus antarctica. Moraines appear to be first colonized by the evergreen N. betuloides, followed within 5–7 years by deciduous N. antarctica. Nothofagus antarctica may replace the former species and develop monospecific stands on glacial valleys. Most trees in the N. antarctica stand studied were older than 40 years and floods may cause a significant mortality of young trees. Recruitment from seed seems to be infrequent. Old-growth stands dominated by deciduous Nothofagus pumilio occupy more stable substrates, and probably represent the last stage of postglacial succession. This long-lived tree species had recorded ages over 200 years. The canopy of N. pumilio forests appears to be a mosaic of even-aged, old-growth patches. We propose that regeneration episodes follow the blowdown of a large portion of the canopy, with long intervals with little or no regeneration. Windstorms may be an important force influencing the regeneration of N. pumilio. Exogenous disturbances, such as floods and windstorms, are an integral part of the forest cycle in the Patagonian region.