Fire history of Araucaria–Nothofagus forests in Villarrica National Park, Chile

Aim In this study we examine fire history (i.e. c. 500 yr bp to present) of Araucaria–Nothofagus forests in the Andes cordillera of Chile. This is the first fire history developed from tree rings for an Araucaria–Nothofagus forest landscape. Location The fire history was determined for the Quillelhue watershed on the north side of Lanin volcano in Villarrica National Park, Chile. The long‐lived Araucaria araucana was commonly associated with Nothofagus pumilio and N. antarctica in more mesic and drier sites respectively. Methods Based on a combination of fire‐scar proxy records and forest stand ages, we reconstructed fire frequency, severity, and the spatial extent of burned areas for an c. 4000 ha study area. We used a composite fire chronology for the purpose of determining centennial‐scale changes in fire regimes and comparing the pre‐settlement (pre‐1883) and post‐settlement fire regimes. In addition, we contrasted Araucaria and Nothofagus species as fire‐scar recorders. Results In the study area, we dated a total of 144 fire‐scarred trees, representing 46 fire years from ad 1446 to the present. For the period from ad 1696 to 2000, using fire dates from Araucaria and Nothofagus species, the composite mean fire interval varied from 7 years for all fires to 62 years for widespread events (i.e. years in which ≥ 25% of recorder trees were scarred). Sensitivity to fire was different for Araucaria and Nothofagus species. More than 98% of the fires recorded by Nothofagus species occurred during the 1900s. The lack of evidence for older fire dates (pre‐1900) in Nothofagus species was due to their shorter longevity and greater susceptibility to being killed by more severe fires. Whereas the thin‐barked N. pumilio and N. antarctica are often destroyed in catastrophic fire events, large and thick‐barked Araucaria trees typically survive. The spatial extent of fires ranged from small patchy events to those that burned more than 40% of the entire landscape (c. > 1500 ha). Main conclusions Fire is the most important disturbance shaping the Araucaria–Nothofagus landscape in the Araucarian region. The forest landscape has been shaped by a mixed‐severity fire regime that includes surface and crown fires. High‐severity widespread events were relatively infrequent (e.g. 1827, 1909 and 1944) and primarily affected tall Araucaria–N. pumilio forests and woodlands dominated by Araucaria–N. antarctica. Although there is abundant evidence of the impact of Euro‐Chilean settlers on the area, the relative influence of this settlement on the temporal pattern of fire could only be tentatively established due to the relatively small number of pre‐1900 fire dates. An apparent increase in fire occurrence is evident in the fire record during Euro‐Chilean settlement (post‐1880s) compared with the Native American era, but it may also be the result of the destruction of evidence of older fires by more recent stand‐devastating fires (e.g. 1909 and 1944). Overall, the severe and widespread fires that burned in Araucaria–Nothofagus forests of this region in 2002, previously interpreted as an ecological novelty, are within the range of the historic fire regimes that have shaped this forested landscape.     

Effects of natural disturbance on the regeneration of riparian forests in a Chichibu Mountains,central Japan

Disturbances in the riparian area had a large effect on each stage of regeneration in riparian forests dominated by Fraxinus platypoda in the Chichibu Mountains, central Japan. F. platypoda adapted well to various disturbances in frequency and size such as landslides and canopy gap formation.The spatial distribution and age structure of F. platypoda were studied in relation to the disturbance regime of the riparian zone. The bell-shaped size and age distributions of F. platypoda suggest that F. platypoda trees were established synchronously in a large disturbance site caused by an earthquake landslide about 200 years ago. For the past 200 years, the topography has been stable and the canopy gap has been recovered by advance regenerated saplings.The distribution of F. platypoda saplings was restricted to an abandoned channel and part of a floodplain. These topographical sites formed by gravel provided safe sites for saplings because stream disturbances did not occur for a long time. Channel bars were under low shade stress because of the lack of herbs and a litter layer, which represents a safe site for seedling establishment. However, the seedling bank could have been destroyed by high frequent flooding caused by large typhoons and the establishment of seedlings might have been prevented for a long time.The regeneration process of F. platypoda was explicated based on the gap dynamics theory during the stable period of topography. On the other hand, an even-aged forest was established in a large scale disturbance site.Nomenclature: Ohwi & Kitagawa (1992)     

A long‐term record of Nothofagus dominance in the southern Andes,Chile

Abstract The general model of regeneration dynamics in Nothofagus forests of southern South America could have value in community ecology if predictive relationships between disturbance history, functional traits and site attributes could be identified. Examined here is the proposal that on favourable sites shade‐intolerant Nothofagus are likely not to survive in competition with shade‐tolerant, broad‐leaved evergreen taxa of temperate rain forests, and persistence, thus, is dependent on periodic coarse‐scale disturbance. Comparison of stand dynamics of three old‐growth Nothofagus forests at different elevations in the southern Andes, Chile where deciduous Nothofagus alpina dominates the upper canopy, and examination of the life history trade‐offs of this variation were made. Stem density of all stems ≥5.0 cm d.b.h. was 233–303 stems per hectare, and basal area was 123.9–171.0 m²ha^?1. Maximum lifespan of N. alpina was found to be greater than ca 640 years, exceeding all previously reported ages for this species in the region. Forests had a stable canopy composition for this long‐term, but some appeared to lack effective regeneration of N. alpina in recent years. Regeneration of N. alpina was generally greater in disturbed stands and higher elevation than in undisturbed stands and at lower elevation. Recruitment emerged to be strongly affected by competitive over‐ and understorey associates. There was a gradient of increasing dependence of N. alpina on disturbance towards the more productive end of the environment gradients, and hence less dependence of N. alpina on disturbance for its regeneration towards higher elevation. The study confirms that changes in forest composition may be explained by processes occurring in accordance with the predictions of the existing model of Nothofagus regeneration dynamics, providing stronger evidence specifically directed at mid‐tolerant N. alpina, and by factoring out regeneration dynamics on favourable sites. Thus, for N. alpina, trait differences probably contribute to the competitive advantage over its associates in productive habitats, and may be linked to small‐to‐intermediate‐sized disturbances which inevitably occur as older trees die, enabling N. alpina to persist in forests and therefore maintain species coexistence for the long‐term.     

Tree age,disturbance history,and carbon stocks and fluxes in subalpine Rocky Mountain forests

Forest carbon stocks and fluxes vary with forest age, and relationships with forest age are often used to estimate fluxes for regional or national carbon inventories. Two methods are commonly used to estimate forest age: observed tree age or time since a known disturbance. To clarify the relationships between tree age, time since disturbance and forest carbon storage and cycling, we examined stands of known disturbance history in three landscapes of the southern Rocky Mountains. Our objectives were to assess the similarity between carbon stocks and fluxes for these three landscapes that differed in climate and disturbance history, characterize the relationship between observed tree age and time since disturbance and quantify the predictive capability of tree age or time since disturbance on carbon stocks and fluxes. Carbon pools and fluxes were remarkably similar across the three landscapes, despite differences in elevation, climate, species composition, disturbance history, and forest age. Observed tree age was a poor predictor of time since disturbance. Maximum tree age overestimated time since disturbance for young forests and underestimated it for older forests. Carbon pools and fluxes were related to both tree age and disturbance history, but the relationships differed between these two predictors and were generally less variable for pools than for fluxes. Using tree age in a relationship developed with time since disturbance or vice versa increases errors in estimates of carbon stocks or fluxes. Little change in most carbon stocks and fluxes occurs after the first 100 years following stand‐replacing disturbance, simplifying landscape scale estimates. We conclude that subalpine forests in the Central Rocky Mountains can be treated as a single forest type for the purpose of assessment and modeling of carbon, and that the critical period for change in carbon is < 100 years.     

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.     

Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

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.     

The influence of resprouting forest canopy species on richness in Southern Cape forests, South Africa

We investigated the relationship between species richness and numbers and types of individuals and species present in forests with different physiognomies in the southern Cape Province, South Africa. Data were collected from three different ‘plot’ types: 400 m², canopy‐scaled (plot length is directly proportional to canopy height) and per 100 individuals closest to a point. Plots were designed to control for the effect of scale on local richness. Canopy species richness was inversely proportional to the abundance of resprouting species. The strength of the relationship between the abundance of resprouters and canopy species richness increased progressively from the 400 m² plots to the canopy‐scaled plots and finally to the plots of 100 individuals. Resprouter abundance decreased, while canopy species richness increased, with increasing canopy height. Resprouters are able to retain their in situ position in the forests for longer periods of time than do reseeders. This reduces individual and species turnover, thus reducing species richness in resprouter‐dominated forests.     

人为干扰对风水林群落林下木本植物组成和多样性的影响

华南地区的风水林是乡村聚落的一种特色林分, 具有守护村庄的象征意义。在过去的数百年中, 风水林在乡村的社会文化习俗的影响下而受到保护, 对当地的生物多样性保育有着重要作用。为揭示人为干扰对风水林的影响, 我们选择广东省东莞市大岭山镇同一林分起源的3个具有相似地形的风水林, 研究了在不同干扰强度下其林下木本植物种类组成和物种多样性。多响应置换过程(multi-response permutation procedures, MRPP)分析表明, 人为干扰显著改变了风水林林下木本植物组成(P = 0.001, A = 0.3886), 沿着干扰由弱至强的梯度呈现出中生性植物减少、阳生性植物递增的趋势。多样性指数变化趋势为重度干扰＞中度干扰＞轻度干扰, 但没有表现出统计学意义上的差异(P＞0.05)。随着干扰强度的增大, 3个风水林群落相互间的林下物种相似性降低, 物种替代率呈增加趋势。双向聚类分析较好地反映出林下物种因受不同人为干扰强度影响而表现出在空间分布上的差异。指示种分析进一步确定了不同干扰强度下具有显著指示值(IV ≥60)的指示种。综合分析表明, 人为干扰有利于阳性物种在风水林内定居生长, 并明显地改变了林下木本植物组成, 但未能引起物种多样性的显著差异。此外, 找出对人为干扰产生关键生态响应的林下指示种, 对增进风水林的生物多样性保育以及生态系统管理有着重要的理论意义和实践价值。     

兴安落叶松天然林碳储量及其碳库分配特征

兴安落叶松天然林作为大兴安岭林区的主要植被类型,在森林生态系统碳循环中具有重要的作用。在大兴安岭林区选择不同林龄的兴安落叶松天然林,调查其乔木、灌草、枯落物和土壤,并结合已建立的单木异速生长方程分别计算其碳储量,以期为明确该地区碳库动态及其碳库分配特征提供参考。结果表明,兴安落叶松天然林总碳储量随林龄的增加逐渐增大,由幼龄林到过熟林分别为140.46、186.63、208.64、308.62和341.03 Mg C/hm2,整体表现为碳汇,这主要与乔木碳储量随林龄的增加逐渐增大有关;乔木碳库的变化范围为45.44—212.67 Mg C/hm2,且其占总碳储量的比例也随林龄的增加逐渐增大,由幼龄林的32.60%到过熟林的62.36%;灌草碳储量占总碳储量的比例较小,仅为0.48%—0.93%;枯落物碳库在过熟林中较多,为26.11Mg C/hm2,这与过熟林较少的人为干扰有关;土壤碳储量以幼龄林最小,成熟林最高,分别为78.06和131.93 Mg C/hm2,但这与我国其他地区天然林相比均较低,这与该地区较浅的土壤发生层有关;土壤碳储量随林龄的变化并不明显,但其占总碳储量的比例却随林龄的增加逐渐减小,由幼龄林的56.01%减小到过熟林的29.35%。     

Impacts of selective logging on population structure and dynamics of a canopy tree (Olea capensis) in Kakamega forest

This study assessed the impacts of logging on the population structure and dynamics of Elgon teak (Olea capensis), a canopy dominant in Kakamega forest. The objective was to ascertain and determine the regeneration status of this tree species in view of its commercial value. A comparison was made between logged and unlogged sections at three sites in the forest. Pairwise comparisons of the differences in the population structure among three sites and the differences in the mean tree size revealed significant differences in all three study sites. Results also revealed a variation in stem density that was dependent on the block of forest under study. Results of the spatial contagion revealed that O. capensis had a varied clumped distribution in all the study sites. Clumping was highest in the Kisere forest. These results point to logging as the major factor that explains the differences in population structure and stem density in different parts of Kakamega forest. And while intermittent recruitment may explain clumped distribution, logging appears to explain the variation in the degree of clumping.     

Catastrophic disturbance, logging and the ecology of mahogany (Swietenia macrophylla King): grounds for listing a major tropical timber species in CITES

Recent debate on whether or not mahogany ( Swietenia macrophylla King) is threatened by the international timber trade has focused on the breadth of its range and estimates of the remaining stock of mahogany trees. These data are inadequate to reveal the status of mahogany populations, both because they are incomplete in areal extent and because they do not reveal population parameters such as the existence or density of young trees smaller than commercial size. However, there is sufficient information on the regeneration ecology of mahogany to indicate that under natural conditions this species regenerates in essentially even-aged stands after catastrophic disturbances destroy many or most trees, and, in the case of fires and flooding, saplings and seedlings as well. Adult mahoganies tend to survive these events, and regenerate by shedding seed onto the resulting gaps or clearings. This ecological strategy makes mahogany vulnerable to logging, first because juvenile mahoganies are not found in the understorey, and secondly because logging operations shortcircuit mahogany regeneration processes by selectively removing almost all mahogany seed sources while leaving standing competing vegetation of other species. Listing of mahogany in CITES Appendix II could provide both a mechanism to fill in gaps in information and an incentive to change current practices in favour of silvicultural management to provide for regeneration of this valuable timber species in forests subjected to logging.     

Effects of Species, Density, Patch-type, and Season on Post-dispersal Seed Predation in a Puerto Rican Pasture1

An experimental study of seed removal of four woody species in an abandoned pasture revealed significant main effects of species, density (higher densities survived more than lower densities), and patch‐type (seeds under shrub patches survived more than in grass patches), but no effects of season. Rates of seed loss decreased with seed size across species. Significant interactions between species and density and between species and patch‐type were also observed.