Regional tectonics, disturbance, and population size of isolated stands of Nothofagus fusca (Nothofagaceae) in a forest ecotone in south-western New Zealand

Aim New Zealand's cool temperate forests are usually dominated by one or more of the five native taxa of Nothofagus (Nothofagaceae; southern beech), but in certain regions there are sharp boundaries against podocarp–broadleaved forest where Nothofagus is rare or completely absent, either for historical (Pleistocene Glaciation) or climatic/biological (mild superhumid climate and competition) reasons. The dynamics of a Nothofagus boundary was investigated by monitoring disturbance-initiated establishment of isolated stands of N. fusca at the extreme limits of its regional distribution. Location The research was carried out in a regional forest ecotone between Nothofagus forest and podocarp–broadleaved forest in the upper Taramakau Valley, South Island, New Zealand. The survey region straddles a major, active fault system and associated tectonic movements and earthquakes with more distant epicentres have contributed to intermittent canopy disturbance of the local forests. Methods Isolated stands of Nothofagus fusca beyond the limits of continuous Nothofagus forest were investigated during two field surveys, separated by 7–10 years. Changes in population size, stem diameter of individual trees, stand basal area and mean annual diameter increment were calculated for each of fifty-four isolated stands. Types of past and recent disturbance and the probable cause of mortality of trees were noted. Results The total population of fifty-four sample stands, ranging in size from one to > 400 stems, increased by 37.4%, and compound basal area increased by 4.7% between the two surveys. Mean stem diameter growth of isolated stands was lower than expected by empirical data for N. fusca, suggesting reduced wood increment at the limits of its distribution. Tree mortality was 0.8% per year. Fifty-one per cent of the dead stems had died as a consequence of various types of natural disturbance, uproots being more common than snaps and crown breakage. Main conclusions The isolated N. fusca stands preferentially occupy sites likely to experience intermittent disturbance, mostly including disturbance of the soil cover, which facilitates their initial establishment and persistence. Because of causal relationships between mass movement on steep slopes and erosion/deposition of talus fans and river terraces, disturbance-initiated changes in forest composition are observed across a range of different landforms.     

Structure and composition of riparian forests with special reference to geomorphic site conditions along the Tokachi River,northern Japan

The structure and composition of riparian forests were examined along the Tokachi River, northern Japan. Both the hydrogeomorphic gradient and the temporal gradient were analyzed in attempt to explain the present pattern of riparian forests. The stability of floodplain surfaces was estimated on the basis of the elevation above the riverbed and the distance from the river channel. The characteristics of the substratum on which trees were established were also examined by excavation of buried sediment. The results indicated that soil moisture and organic content increased while the size of particles in the substratum decreased with increased elevation and distance from the river channel. Gradient analysis was employed to examine the distribution of dominant species, such as Alnus hirsuta, Toisusu urbaniana, Populus maximowiczii, Picea jezoensis and Abies sachalinensis. The relative dominance of each could be arrayed across the elevation gradient. Although the three broad-leaved pioneers dominated bars and floodplains near the river channel, their modes shifted from lower to higher elevation and amplitudes of distribution curves decreased in the following order: A. hirsuta, T. urbaniana, P. maximowiczii and conifers, which were located on the highest floodplains. Sites could be divided into three classes in terms of stability. There were fewer species at active sites, which favored the three pioneer species, but species richness and diversity increased with stand age. Semi-active and stable sites were more diverse with the establishment of conifers and other broad-leaved trees, which included upland species. However, species richness peaked and then decreased after trees reached 50 to 60 years of age. The growth of dwarf bamboo and the development of conifer-dominant stands impeded the establishment of other species, thereby reducing species richness and diversity in mature stands. Chronologically, floodplains could be differentiated into high- and low-frequency zones of flood disturbance, with pioneer species occupying the former, and late successional species found largely in the latter.     

Pollen flow in fragmented landscapes maintains genetic diversity following stand-replacing disturbance in a neotropical pioneer tree,Vochysia ferruginea Mart

In forests with gap disturbance regimes, pioneer tree regeneration is typically abundant following stand-replacing disturbances, whether natural or anthropogenic. Differences in pioneer tree density linked to disturbance regime can influence pollinator behaviour and impact on mating patterns and genetic diversity of pioneer populations. Such mating pattern shifts can manifest as higher selfing rates and lower pollen diversity in old growth forest populations. In secondary forest, where more closely related pollen donors occur, an increase in biparental inbreeding is a potential problem. Here, we investigate the consequences of secondary forest colonisation on the mating patterns and genetic diversity of open-pollinated progeny arrays for the long-lived, self-compatible pioneer tree, Vochysia ferruginea, at two Costa Rican sites. Five microsatellite loci were screened across adult and seed cohorts from old growth forest with lower density, secondary forest with higher density, and isolated individual trees in pasture. Progeny from both old growth and secondary forest contexts were predominantly outcrossed (t_m=1.00) and experienced low levels of biparental inbreeding (t_m−t_s=0.00–0.04). In contrast to predictions, our results indicated that the mating patterns of V. ferruginea are relatively robust to density differences between old growth and secondary forest stands. In addition, we observed that pollen-mediated gene flow possibly maintained the genetic diversity of open-pollinated progeny arrays in stands of secondary forest adults. As part of a natural resource management strategy, we suggest that primary forest remnants should be prioritised for conservation to promote restoration of genetic diversity during forest regeneration.     

Simulation modeling of the effects of site conditions and disturbance history on a boreal forest landscape

Abstract. Computer simulations were used to elaborate hypotheses about controls on forest structure and composition in a 0.7 km ≤ area of boreal forest in Central Sweden. DBH and species of all adult trees and stand conditions were recorded for 57–10 m radius plots. Ordination of these data suggested that nutrient-availability and time-since-disturbance were the main controls of forest composition and structure within the area. The simulation model couples equations representing the effect of tree canopy structure and biomass on light and soil conditions with equations representing the effect of these conditions on reproduction, growth and mortality in height cohorts of trees on a 0.1 ha patch. Nitrogen-availability levels for each modeled plot were simulated by species-specific growth multipliers. The model was run for 400 simulated yr at six levels of N availability. Age and N status of each study plot were inferred by matching with the most similar model output. Inferred ages agreed with what is known of the disturbance history, and site factors related to soil fertility were correctly correlated with the inferred N status. The consequences of size-selective disturbance were explored by model experiments. Biomass was removed from large or small size classes at 100 - 200 yr and the simulations were run for an additional 300 yr. Disturbed stands of high N status often became similar to undisturbed stands of different N status. Size-selective disturbances produced stands that were different from any in the undisturbed succession, but these differences disappeared within 50 - 100 yr, implying successional convergence in stand structure and composition. Plots of simulated basal area against time and nitrogen-availability for the four species illustrate the time dependence of species performance along a fertility gradient.     

Impacts of forest fragmentation on the mating system and genetic diversity of white spruce (Picea glauca) at the landscape level

We studied the mating system of white spruce (Picea glauca) in a landscape fragmented by agriculture in northern Ontario, Canada. We sampled 23 stands that ranged in size from 1 to >500 trees isolated by 250-3000 m from the nearest other stand. Six polymorphic allozyme loci from four enzyme systems were used to genotype approximately 10 000 embryos from 104 families. We detected no allele frequency heterogeneity in the pollen pool among stands or families (Phi(FT)=-0.025). Overall, estimates of outcrossing were high (t(m)=94% and mean t(s)=91%) but significantly different from unity. Bi-parental inbreeding (t(m)-t(s)=3.2%) was low but significantly different from zero. Allozyme-based outcrossing estimates did not differ significantly among three stand-size classes (SSCs): small (<10 trees), medium (10-100 trees) and large (> or =100 trees). The number of effective pollen donors was high in all SSCs, but was significantly lower in small stands (N(ep)=62.5) than in medium-sized and large stands (both N(ep)=143). The primary selfing rate was significantly higher in medium stands than in large stands. We found no significant difference in genetic diversity measures in the filial (seed) population among SSCs. Overall, these results indicate that white spruce stands in this fragmented landscape are resistant to genetic diversity losses, primarily through high pollen-mediated gene-flow and early selection against inbred embryos. We discuss the importance of using seed data, in conjunction with genetic data, to evaluate the impacts of fragmentation on natural populations.     

Regeneration dynamics and life history differences in southern Chilean Nothofagus forests: a synthesis

Large, infrequent natural disturbances have been proposed as a key component in determining the distribution and abundance of Nothofagus in southern Andean temperate forests. In this study, a comparison of influences of small-to-intermediate-sized natural disturbances and effects of selective logging on the establishment and growth of Nothofagus forests in south-central Chile are synthesized with the results of other forest studies to develop general hypotheses on the regeneration dynamics of southern Chilean Nothofagus alpina forests. A synthesis is given for regeneration pattern in the abundance of 18 temperate rain forests in the Andean Range, Chile. The study aimed to determine the role of life history differences in promoting coexistence of the five main tree species (N. alpina, N. dombeyi, N. pumilio, N. macrocarpa and Laurelia philippiana) in N. alpina-dominated forests. Age data reported for N. alpina, a shade mid-tolerant emergent tree in the temperate rain forests of southern Chile, indicate maximum lifespans > 650 years, figures unprecedented for N. alpina. In low elevation stands, N. alpina coexists with broad-leaved evergreen tree species, such as L. philippiana, L. sempervirens, Persea lingue, and in these stands an intermittent establishment of Nothofagus occurred and appeared to be most dependent on small-to-intermediate disturbances. In high-elevation stands, in contrast, Nothofagus establishment was less dependent on disturbance, regeneration being much more continuous even in the absence of canopy openings. The forests studied provide another example of the general pattern of increasing dependence of Nothofagus on disturbance towards the more productive end of the environmental gradients. As a long-lived pioneer and despite its dependence on disturbance in lowland sites, N. alpina has been subject to selection for complementary growth, adult survivorship, and mid-tolerance to shade. Thus, interspecific differences in juvenile and adult life history characteristics of N. alpina and its competitors may be sufficient to maintain its persistence in the landscape. In conclusion, this study of population structures and replacement patterns provides a comprehensive picture on our understanding of regeneration dynamics and trait differentiation in southern temperate forests by recognizing both the influences of environmental gradients (i.e., altitude) on competing species and the disturbance regimes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Earthquake impacts in old‐growth Nothofagus forests in New Zealand

Abstract. Six stands located on different land forms in mixed old‐growth Nothofagus forests in the Matiri Valley (northwest of South Island, New Zealand) were sampled to examine the effects of two recent large earthquakes on tree establishment and tree‐ring growth, and how these varied across land forms. 50 trees were cored in each stand to determine age structure and the cores were cross‐dated to precisely date unusual periods of radial growth. The 1968 earthquake (M = 7.1, epicentre 35 km from the study area) had no discernible impact on the sampled stands. The impact of the 1929 earthquake (M = 7.7, epicentre 20 km from the study area) varied between stands, depending on whether or not they had been damaged by soil or rock movement. In all stands, the age structures showed a pulse of N. fusca establishment following the 1929 earthquake, with this species dominating establishment in large gaps created by landslides. Smaller gaps, created by branch or tree death, were closed by both N. fusca and N. menziesii. The long period of releases (1929–1945) indicates that direct earthquake damage was not the only cause of tree death, and that many trees died subsequently most likely of pathogen attack or a drought in the early 1930s. The impacts of the 1929 earthquake are compared to a storm in 1905 and a drought in 1974–1978 which also affected forests in the region. Our results confirm that earthquakes are an important factor driving forest dynamics in this tectonically active region, and that the diversity of earthquake impacts is a major source of heterogeneity in forest structure and regeneration.     

Influence of Invasive Tree Kill Rates on Native and Invasive Plant Establishment in a Hawaiian Forest

Fire tree (Morella faya) has invaded extensive areas of wet and mesic forest on the Island of Hawai’i, forming nearly monospecific stands. Our objective was to identify a method of controlling M. faya, which would allow native plants to establish while minimizing establishment by invasive plants. Treatments (logging all trees, trees left standing but girdled, and incremental girdling over 20 months) were selected to kill M. faya stands at different rates, thereby creating different conditions for species establishment. Leaf litter was either removed or left in place; seeds and seedlings of three native pioneer species, three native forest species, and three alien invasive species were then added to determine their ability to establish. Native pioneer species established best in the log and girdle treatments, whereas seedling emergence of native forest species was higher in the girdle and incremental girdle treatments. Seedlings of invasive species emerged faster than the natives, but each of them responded differently to the stand treatments. Leaf litter reduced seedling emergence for all species, with small‐seeded species (<1 mg/seed) affected most under low light conditions. No single method eliminated all invaders, but girdling of M. faya provided suitable conditions for most native species. If combined with selective removal of the most disruptive alien species and native seed additions, girdling could be an effective general strategy for restoring native forests that have been overwhelmed by woody invaders.     

不同干扰强度下针阔混交林树种空间格局及种间关联性

为了探究不同干扰程度下针阔混交林树种空间格局变化及树种种间关联性,本研究以关帝山庞泉沟自然保护区不同干扰强度的针阔混交林为对象,选取郁闭度、林分密度、伐桩数量作为划分干扰程度的因子,采用相对影响法,将林分划分为未干扰、中度干扰和严重干扰3个等级.利用空间点格局K2函数分析了林分的空间分布格局及其不同树种的种间关联.结果...     

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.     

The size and composition of soil seed-banks in remnant patches of three structural rainforest types in North Queensland

The size and species composition of the soil seed-bank in a remnant patch of each of three structurally and floristically distinct rainforests (Complex Mesophyll Vine Forest, Complex Notophyll Vine Forest and Semi-Evergreen Vine Thicket) were assessed. Seeds of 94 species germinated from 12 surface soil samples collected from each site. All three seed-banks were composed mostly of herbs characteristic of roadsides and agricultural land, and pioneer rainforest trees and shrubs. Agglomerative classifications indicated that the seed-bank samples from each rainforest remnant had a characteristic species composition and could be distinguished reliably from seed-bank samples drawn from other sites. Seeds of species present in the standing forest were poorly represented in the seed-banks except for one long-lived pioneer tree, Dendrocnide photinophylla, at one site. The seed-bank from the seasonally dry vine thicket was significantly larger (4000 seeds m^-2) than those from the two moister sites (400–600 seeds m^-2, contained more seeds of roadside and agricultural herbs, and fewer seeds of rainforest pioneer and secondary shrubs and trees. We suggest three explanations for the different seed-bank structure observed in the seasonally dry forest site. First, with increased deciduousness in rainforests, seed-banks are increasingly subject to invasion and domination by seeds of rapidly maturing herbs. Second, long-lived seeds that germinate in canopy gaps would be less likely to accumulate under deciduous forests because they would he exposed annually to conditions suitable for germination. Third, chronic disturbance by cattle and pigs produces sites suitable for the establishment of rapidly maturing herbs, and possibly disperses their seeds into the forest.     

Roles of Birds and Bats in Early Tropical-Forest Restoration

Restoration of tropical forest depended in large part on seed dispersal by fruit-eating animals that transported seeds into planted forest patches. We tested effectiveness of dispersal agents as revealed by established recruits of tree and shrub species that bore seeds dispersed by birds, bats, or both. We documented restoration of dispersal processes over the first 76 months of experimental restoration in southern Mexico. Mixed-model repeated-measures randomized-block ANOVAs of seedlings recruited into experimental controls and mixed-species plantings from late-secondary and mature forest indicated that bats and birds played different roles in the first years of a restoration process. Bats dispersed pioneer tree and shrub species to slowly regenerating grassy areas, while birds mediated recruitment of later-successional species into planted stands of trees and to a lesser extent into controls. Of species of pioneer trees and shrubs established in plots, seven were primarily dispersed by birds, three by bats and four by both birds and bats. Of later-successional species recruited past the seedling stage, 13 were of species primarily dispersed by birds, and six were of species dispersed by both birds and bats. No later-successional species primarily dispersed by bats established in control or planted plots. Establishment of recruited seedlings was ten-fold higher under cover of planted trees than in grassy controls. Even pre-reproductive trees drew fruit-eating birds and the seeds that they carried from nearby forest, and provided conditions for establishment of shade-tolerant tree species. Overall, after 76 months of cattle exclusion, 94% of the recruited shrubs and trees in experimental plots were of species that we did not plant.     

The Effects of Land-use History on Soil Properties and Nutrient Dynamics in Northern Hardwood Forests of the Adirondack Mountains

Soil nutrient pools and nitrogen dynamics in old-growth forests were compared with selectively logged stands and stands that were selectively logged and then burned approximately 100 years ago to test the hypothesis that land-use history exerts persistent controls on nutrient capital and nitrogen (N) transformation rates. We provide estimates of net N mineralization and nitrification rates for old-growth forests from the northeastern United States, a region in which few old-growth forests remain and for which few published accounts of mineralization rates exist. At the plot level, no effects of the dominant tree species were observed on any measured soil properties or N-cycling rates. Effects of alternate disturbance histories were detected in soil carbon (C) and N pools. Old-growth forest soils had higher total C (67 Mg·ha^–1) and N capital (3.3 Mg·ha^–1) than that of historically logged then burned soils (C = 50 Mg·ha^–1 and N = Mg·ha^–1), with intermediate values (C = 54 Mg·ha^–1 and N = 2.7 Mg·ha^–1) in the stands that were historically logged. Despite these differences in C and N content, corresponding differences in C–N ratio, net N mineralization rates, and net nitrification rates were not observed. The N concentration in the green foliage of American beech trees (Fagus grandifolia) was also highest from canopy trees growing in old-growth stands (3.0%), followed by logged stands (2.6%), and lowest in the logged/burned stands (2.2%). These data suggest that some legacies of light harvesting on ecosystem processes may be detected nearly 100 years following the disturbance event. These results are discussed in the context of how multiple forest disturbances act in concert to affect forest dynamics.     

The spatial characteristics of stand structure in Pinus torreyana

The arrangement of trees within a stand by location and age (stand structure) is in part determined by the life history strategy of the species and the disturbance history of the stand. In western North America such disturbances are often the product of wildfires and human management activities. The current study uses spatial analysis to characterize three stands of Pinus torreyana with known disturbance histories. Two stands are located at Torrey Pines State Reserve (TPSR). Of these, one stand has burned twice since 1972. Fire has been successfully excluded from the other stand since the early part of this century. A third stand, on Santa Rosa Island, Channel Islands National Park (SRI), has been grazed heavily since the mid-19th century and has not experienced fire since that time. One-hectare study plots were established in the interior of each stand. Considering the known life history attributes of P. torreyana, and the disturbance histories of the stands, predictions are made concerning the spatial characteristics of the respective stands. All trees within each study plot were sized by diameter breast height (DBH) and mapped. Three techniques of spatial analysis are applied to the resulting unweighted point pattern distributions and the distributions weighted by the square root of DBH as a surrogate for age. The results are consistent with predictions and confirm the following generalizations concerning patterns of aggregation in Torrey pine stands. Young trees tend to be more aggregated than old trees within the same stands. Young stands tend to be more aggregated than old stands on otherwise ecologically similar sites. On a periodically disturbed site there are clusters of trees that represent cohorts of post-disturbance recruitment.     

Releasing Rain Forest Succession: A Case Study in the Dicranopteris linearis Fernlands of Sri Lanka

Throughout tropical moist climates, Dicranopteris linearis fernlands can develop as a result of rain forest clearance followed by frequent burning. In Sri Lanka, D. linearis fern-lands are capable of suppressing the regeneration of rain forest. Field experiments were conducted at Sinharaja Man and Biosphere Reserve, a rain forest where fernlands occupy substantial areas of the reserve boundary. The experiment's objective was to identify methods for initiating forest regeneration in fernlands dominated by D. linearis Three disturbance treatments were used to initiate seedling regeneration: clean weed, root removal, and till. We hypothesized that increasing the severity of the soil disturbance would establish vegetation with higher species richness and diversity, greater above-ground dry biomass, and higher percentage cover and seedling density. Results indicate only partial support for this hypothesis. Dry biomass was greatest in till treatments, the most severe soil disturbance. By comparison, species richness and diversity, seedling density, and percentage cover were greatest in root-removal treatments, though in many instances the differences were not significant. The study clearly demonstrated that any kind of soil disturbance can facilitate the establishment of herbs, shrubs, and trees in a fernland dominated by D. linearis. Results showed that herbs, sedges, grasses, and pioneer shrubs represented greater proportions of seedling recruits than did pioneer trees. Seedlings of primary-forest tree species were nearly nonexistent. In general, results showed that soil disturbance can play an important role in site preparation for the purpose of initiating non-fern vegetation in fernlands dominated by D. linearis.     

Succession: A population process

Summary Recent critical reviews suggest the need for a reductionistic approach to the study of secondary plant succession. We propose viewing succession as the result of the underlying plant population dynamics. This approach is being developed using nearly 50 years of permanent sample plot records.After initial establishment Pinus taeda shows an exponential depletion with stands of various densities conforming to the reciprocal yield relationship. Uneven-aged hardwoods also show exponential depletion. Canopy disturbance can enhance the establishment process, though severe disturbance and the consequent abundant regeneration can lead again to dense, even-aged stands with low levels of establishment. These results suggest a general pattern of forest development wherein establishment is initially important, but is quickly replaced by mortality as the dominant process when the dense, even-sized stand starts to thin. Eventually, failing additional disturbance, natural mortality will again open the canopy allowing development of a balance between establishment, and mortality.This research was supported by National Science Foundation grants DEB-7708743 and DEB-7804043 to R.K.P. and DEB-7707532 and DEB-7804041 to N.L.C.     

The Seed and Fern Spore Bank of a Recovering African Tropical Forest

Seed banks contribute to forest regeneration after disturbance, but less is known about fern spore banks, particularly in a paleotropical context. We sampled the buried seed and fern spore bank in Mabira Forest, a 300 km² forest in central Uganda, to explore the effect of time since disturbance. Soil cores (5 cm depth) were taken from 39 plots across three different classes of ‘recovery’: (1) not disturbed since 1950; (2) logged between 1950 and 1980; and (3) cleared for agriculture between 1970 and 1990 but reforested since. Plant emergence was monitored in a glasshouse. We predicted that the seed bank would reflect time since disturbance, with more pioneer species in recently disturbed stands, and that the fern spore bank would reflect stand age less closely due to greater dispersal capacity. We recorded a median 752 seeds per square meter, most of which were trees; the most abundant species was the invasive tree Broussonetia papyrifera. The fern spore bank was twice as dense, but 95 percent of fern spores were of one species, Christella parasitica. Tree seed density was significantly affected by time since disturbance with fewer seeds in the older stands. Herb seed density, fern spore density, and species richness for all groups were not significantly affected by time since disturbance. Neither seed bank nor fern spore bank closely resembled the aboveground vegetation. We compared our results to existing literature on seed banks in tropical forests, finding that our densities are relatively high for African forests, but low compared to the Neotropics and Australia.     

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

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

Sexual and Asexual Reproduction in Natural Stands of Populus nigra

The balance between sexual and asexual propagation in the establishment of natural stands of Populus nigra was assessed from the genotypes of trees sampled in different natural stands. One site was completely sampled and genotyping proceeded using STS and RAPD markers. Among 118 trees, only four were vegetative copies of other trees of the site. Isozymes were used in four stands, and a method to detect the presence of vegetative copies in a sample is proposed for these markers. No vegetative copies were found in two cases, and a low number of copies was detected in the other two. Observations are reported for seedlings as well as for different modes of vegetative propagation in natural stands. We conclude that sexual and asexual propagation play complementary roles in the dynamics of this species, and that in most cases the adults originate from seedlings.     

Conifer regeneration failure in New Zealand: dynamics of montane Libocedrus bidwillii stands

Size and age structure analysis were used to reconstruct the population dynamics of six apparently old and stable stands dominated by the conifers Libocedrus bidwillii and/or Dacrydium biforme in Camp Creek, Westland, New Zealand. All stands contained relatively even-aged groups of Libocedrus that had probably originated after either mass movement or windthrow. The age distributions of stands reflected the effects of disturbances 200–400 yr ago.There was no evidence to support synchronous failure of regeneration of Libocedrus and Dacrydium biforme. Between 1600 and 1800 A.D., the period of the postulated conifer regeneration gap, over 70% of trees of these species had established. A lack of Libocedrus regeneration over the last 200 yr reflects a lack of disturbance over that time. Because of the dominating influence of disturbances and other factors it may be inappropriate to use tree population structures alone to substantiate or refute claims of recent climate change.