Historical fire and vegetation dynamics in dry forests of the interior Pacific Northwest,USA, and relationships to Northern Spotted Owl (Strix occidentalis caurina) habitat conservation

Regional conservation planning frequently relies on general assumptions about historical disturbance regimes to inform decisions about landscape restoration, reserve allocations, and landscape management. Spatially explicit simulations of landscape dynamics provide quantitative estimates of landscape structure and allow for the testing of alternative scenarios. We used a landscape fire succession model to estimate the historical range of variability of vegetation and fire in a dry forest landscape (size ca. 7900 km²) where the present-day risk of high severity fire threatens the persistence of older closed canopy forest which may serve as Northern Spotted Owl (Strix occidentalis caurina) habitat. Our results indicated that historically, older forest may have comprised the largest percentage of the landscape (∼35%), followed by early successional forest (∼25%), with about 9% of the landscape in a closed canopy older forest condition. The amount and condition of older forest varied by potential vegetation type and land use allocation type. Vegetation successional stages had fine-grained spatial heterogeneity in patch characteristics, with older forest tending to have the largest patch sizes among the successional stages. Increasing fire severities posed a greater risk to Northern Spotted Owl habitat than increasing fire sizes or frequencies under historical fire regimes. Improved understanding of historical landscape-specific fire and vegetation conditions and their variability can assist forest managers to promote landscape resilience and increases of older forest, in dry forests with restricted amounts of habitat for sensitive species.     

北美地区景观情景分析探索技术

自1996年以来, 景观情景分析探索技术在北美地区就被作为景观动态模拟工具, 广泛应用于森林植被恢复与管理过程。其能够模拟森林景观在不同演替阶段、自然干扰和森林经营管理措施下的动态变化情况, 进而评价和预测各模拟因子对森林景观的影响程度。着重介绍了景观情景分析探索技术的应用范围、组成体系、输入数据要求和进行景观情景分析的主要步骤。最后, 通过实例说明该技术体系的具体应用。     

The impact of fire and density-dependent mortality on the spatial patterns of a pine forest in the Hulun Buir sandland,Inner Mongolia,China

Wildland fire, especially surface fire, is one of the major disturbances in forest ecosystems such as the Mongolian pine forest of the Hulun Buir sandland. However, little is known about the impact of fire on the spatial patterns at the stand level and its consequences for successional dynamics. To fill this gap we mapped three plots of Mongolian pine forest comprising sites of the same age that were affected by fire in 2006 and 1994, and a site without fire. We explored the stand structure using diameter at breast height (DBH) distributions and basal area, and used point pattern analysis to quantify the observed spatial patterns. Null models included homogeneous Poisson and Thomas cluster processes for univariate analyses, toroidal shift to test for independence, and random labelling for analyzing mortality. Large stems showed at all three stands a tendency to regularity, whereas small stems were mostly clustered. However, small stems did not become more regular at plots with earlier fire, and only the “overstocked” 2006 pre-fire stand showed negative association between large and small trees. Mortality was non-random, showing clustering of dead (or surviving) trees and clear density dependence where stems with more neighbors had a higher risk to die. Fire killed mostly smaller trees and after prolonged fire-free periods the stand approached the pre-fire patterns structurally and spatially. Our study showed that surface fire causes strong thinning in later succession stages and pushes the forest into maturity. It may also enhance resistance of our forest to more severe fire under the relatively harsh environmental conditions. A novel test statistic for random labeling allowed directly testing for density-dependent mortality and could be widely applied in point pattern analysis.     

Mycorrhizas and secondary succession in aspen-conifer forests: Light limitation differentially affects a dominant early and late successional species

Plant succession and mycorrhizal fungi both play crucial roles in shaping the development of forest ecosystems. However, despite the strong potential for interactions between them, few studies have examined how patterns of forest succession affect mycorrhizal associations that a majority of plant species depend on to alleviate soil resource constraints. Fire suppression in subalpine forests over the last century has changed successional patterns in ways that may have important implications for mycorrhizal associations of forest tree species. To better understand these relationships we conducted a field and greenhouse study in which we examined mycorrhizal infection along gradients of light intensity and soil nutrient availability that develop as aspen becomes seral to conifers under longer fire return intervals. We examined whether ectomycorrhizal associations of quaking aspen (Populus tremuloides), a shade intolerant, early succession species, were more sensitive to light and soil resource limitations than subalpine fir (Abies lasiocarpa), a shade tolerant, late succession species. In the field study, ectomycorrhizal infection of aspen roots was reduced by 50% in conifer dominated stands relative to aspen stands. In contrast, subalpine fir maintained its EM associations regardless of the successional status of the stand. The greenhouse results were consistent with field results and indicated that light limitation was the driving force behind reductions in EM infection of aspen roots in later stages of succession. These results suggest that nutrient limitations constraining early successional species may be exacerbated by losses in EM associations via light limitations created by late successional species. This is one potential mechanism by which climax forest species create a competitive advantage over early successional species and these results suggest that it is likely exacerbated by longer fire return intervals.     

热带次生林经营研究文献分析

热带次生林面积巨大且在热带森林资源中的作用越来越重要,因而拥有巨大的经营潜力和生态功能。次生林演替过程中的主要研究成果如下：土壤贮存的种子对次生林的再生非常重要,光投射的强度增加和温度提高将刺激种子萌芽,而残余植被群落强烈影响次生林种子的散布,食种子动物影响次生林的成林;次生林群落植物种类数量一般随演替的进程而增加;随着演替的进程,次生林内阳生树种趋于衰亡,耐荫树种不断增加;次生林的树种数量能接近原生林水平的时间过程更随着林分种类、过去土地的利用强度和利用种类,以及环境条件的不同而异;在一定环境中,人工林树种通过人工林树种改变光、温度和土壤表层的湿度,能够促进发芽和种子生长,改善林地和林下植被的状况,从而加速次生林演替。大多数食草动物喜欢在次生林中取食,因为次生林中先锋树种很少或没有机械和化学保护,且林内许多树种产生大量可食用的果实;环境和火对次生林演替有一定影响;在早期的演替过程中,更多的生物量分配到资源需求旺盛的组织（如叶和细根）,在演替晚期阶段,更多的生物量则分配到结构组织中（如木质部和根端）。次生林在15至20年内能迅速积累生物量,然后积累逐渐减慢;次生林被砍伐或火烧后,森林土壤的结构被破坏,导致土壤有机质和氮减少。     

林火干扰与森林群落动态研究综述

林火是森林生态系统最常见的干扰类型,在森林群落演替中起着重要的作用。本文回顾了林火对森林群落演替的影响、物种对火干扰的适应、火干扰与森林更新、火干扰与物种和景观多样性、火干扰森林群落动态的研究方法,讨论了未来研究方向。     

Wind disturbances shape old Norway spruce-dominated forest in Bulgaria

Correct knowledge of disturbance ecology is essential for understanding the characteristic behavior of forest ecosystems and for guiding appropriate management strategies. However, the role of natural disturbances in shaping European mountain forest ecosystems has not been adequately studied, possibly because of the perception that the development of most European forests is primarily shaped by human influences and/or fine-scale gap-phase dynamics.In the present study, we investigate the long-term disturbance history of old protected forest dominated by Norway spruce in the Parangalitsa Reserve, Bulgaria. We used aerial photo interpretation and dendroecological methods to reconstruct the history of wind, insect, and fire disturbances across a topographically complex landscape. Over the past 150 years wind has been the most important disturbance agent in this ecosystem and at least 18% of the forested area shows evidence of high-severity blowdowns. Windthrow patches ranged in size from <1 ha to >10 ha (minimum 0.11 ha, mean 0.16 ha, maximum 10 ha). Although small disturbances were much more frequent, few larger blowdowns accounted for most of the disturbed area. Pure coniferous and single-cohort coniferous forest patches were more affected by blowdowns than mixed coniferous-deciduous and multi-cohort coniferous forest patches. Although bark beetle (Ips typhographus) populations were large enough to cause mortality of some live trees, the populations did not grow to epidemic proportions during recent decades. Fire disturbance was of limited importance in the last 200 years and only two patches (4% of the study area) showed evidence of fire.The present research indicates that wind disturbances have been characteristic of these ecosystems at least over the past decades to centuries. Thus, blowdowns appear integral to the normal function and structure of the Picea-dominated mountain forests in the region and such events, in and of themselves, do not represent unhealthy forest conditions or environmental emergencies. Management strategies that aim to maintain these ecosystems within a natural range of variation should incorporate wind disturbances into the management strategy. The frequency and magnitude of future wind disturbances may be considered within the historical framework described in the current study to assess potential effects of climate change on altered disturbance regimes.     

Effects of deforestation on microbial diversity in a Siberian larch(Larix sibirica) stand in Mongolia

For evaluating microbial community changes in a Siberian larch stand disturbed by forest fire or clearcutting,357 clones were randomly selected and sequenced using a culture-independent approach and 16S rRNA sequencing to characterize the bacterial composition and diversity from the different disturbed Siberian larch stands.Interestingly,the burned larch stand had an increase in the relative amounts of b-proteobacteria and Firmicutes and a decrease in Acidobacteria,while Gemmatimonadetes increased Verrucomicrobia decreased in the harvested larch stand.Microbial diversity and richness were higher in the undisturbed larch stand than the disturbed(burned or clear-cut)larch stands,and the influence of clear-cutting was more negative than that of the forest fire.This study indicates that evaluating the microbial diversity of undisturbed,burned,and clear-cutting Siberian larch stands provides information about the impact of forest disturbances on soil microbial communities,which may be helpful for understanding and evaluating soil health and devising reafforestation strategies for larch.     

火干扰对天然红松林结构和演替过程的影响

Investigations on charcoal in the soil, fire-scarred trees, stand composition, forest structure as well as regeneration status were carried out in the natural broad-leaved/Korean pine (Pinus koraiensis) forest after fire disturbance at Liangshui Nature Reserve on the mid-north of Xiaoxing‘an Mountains from 1990 to 1992, and the ecological effects of fire disturbance on the formation and succession of this kind of forest were analyzed according to the survey results. The average depth of charcoal in the soil was related to the timing of the fire. According to the characteristic of fire-scarred trees, the dynamic map of the fire behavior was drawn onto the topographic map. It showed that the dimension and extent of the fire disturbance was closely related with site conditions. Fire disturbance only led to a significant difference in stand composition and diameter class structurefor the stands at different locations, rather than completely destroying the forest. After fire disturbance, the horizontal community structure was a mosaic of different patches, which were made up of different deciduous species or different sizes of Korean pines, and the succession trend of each patch was also different. In the sites with the heavy fire disturbance, the intolerant hardwood species were dominant, and there were a large number of regenerative Korean pine saplings under the canopy. In the moderate -disturbed sites, the tolerant hardwood species were dominant, and a small number of large size Korean pines still survived. In the light-disturbed sites, large size Korean pines were dominant.     

From clearfell coupe to old-growth forest: Succession of bird assemblages in Tasmanian lowland wet eucalypt forests

As forests undergo succession after major disturbance events their assemblages of birds also change. Thus the frequency and extent of wildfire or clearfelling in the landscape can potentially affect the species-richness and abundance of forest birds. We used a chronosequence approach to investigate succession of bird communities in Tasmanian lowland wet eucalypt forest, from shortly after disturbance through to old-growth forest aged approximately 200–250 years. The number of native bird species recorded per survey per site increased as a linear function of stand-age. However, succession did not involve a unidirectional transition in assemblage-composition because of differences in successional responses among individual species and also among guilds of birds that mostly inhabited different strata of the forest. This was exemplified by the crescent honeyeater, which was observed most frequently in the youngest (6–8 years) and oldest (200–250 years) forests that we surveyed, and by the superb lyrebird (introduced to Tasmania from mainland south-eastern Australia) which favoured mid-aged regrowth (42–43 years) after clearfelling. Forests aged 200–250 years had the greatest richness-per-survey of those native species that were observed mostly in the lower layer or mid-layer. However, the greatest richness-per-survey of canopy-dwelling species and the highest native species-richness across the survey period were found in forests aged around 150 years. Younger regrowth was generally less rich in birds, although regrowth in the first decade after clearfelling was the only seral stage inhabited by superb fairy-wrens. These results suggest that, in the Tasmanian lowland wet eucalypt forest landscape, species-richness of birds may be greatest when old-growth forest is interspersed with young regrowth forests. Hence for bird conservation, a challenge is to ensure that old-growth forest continues to prevail in the production forest landscape.     

Recovery of understory vegetation biomass and biodiversity in burned larch boreal forests in Northeastern China

Recovery of biomass and biodiversity of forest understory vegetation after fire disturbance has been widely studied; however, how this relationship changes and what are the determinants at different post-fire stages in larch boreal forests are still unclear. We investigated a chronosequence of 81 understory plots in larch boreal forests that were disturbed by fires in 1987 (S5), 1992 (S4), 1996 (S3), 2002 (S2), or 2007 (S1). Analysis of variance was conducted to test the differences of biodiversity and biomass among various post-fire stages. Different regression models were used to fit the relationship between biomass and biodiversity, while factors influencing this relationship were identified by boosted regression tree analysis. Results showed that total understory biomass increased from 2.51?t?ha^?1 in S1 to 8.47?t?ha^?1 in S3 and declined to 5?t?ha^?1 in S5. Similar dynamics were also found between species richness and species diversity. Positive linear correlations linked biomass and biodiversity throughout most of the post-fire periods. Slope and stand density were the two most important factors influencing the secondary succession of understory vegetation after fire. Geographical factors and overstory competition determine the orientation of vegetation recovery, and the impacts of climate on vegetation are muted after fire disturbances.     

林火与病虫害的相互作用

火是森林生态系统演替过程中的一种自然干扰因子, 在促进森林发育、维持生物多样性特别是景观多样性方面起着积极的作用。森林中的多种干扰间有关联性, 火灾、虫害和病原体之间存在着复杂的相互作用。文中综述了火干扰与森林病虫害的相互影响。火灾会引起森林组成和景观的变化, 进而影响森林病虫的生存与发生条件。病虫害通过改变可燃物特性及其分布, 影响火烧频度和强度。这一研究领域的未来发展趋势是更注重火与病虫害之间的量化关系, 研究结果将为开展科学的林火管理提供理论依据, 实现森林经营的可持续。     

Post-fire soil nitrogen content and vegetation composition in Sub-Boreal spruce forests of British Columbia's central interior,Canada

Forest fires are known to influence nutrient cycling, particularly soil nitrogen (N), as well as plant succession in northern forest ecosystems. However, few studies have addressed the dynamics of soil N and its relationship to vegetation composition after fire in these forests. To investigate soil N content and vegetation establishment after wildfire, 13 sites of varying age class were selected in the Sub-Boreal spruce zone of the central interior of British Columbia, Canada. Sites varied in time since the last forest fire and were grouped into three seral age classes: (a) early-seral (<14 years), (b) mid-seral (50–80 years) and (c) late-seral (>140 years). At each site, we estimated the percent cover occupied by trees, shrubs, herbs and mosses. In addition, the soil samples collected from the forest floor and mineral horizons were analyzed for the concentrations of total N, mineralizable N, available NO₃⁻-N and available NH₄⁺-N. Results indicated that soil N in both the forest floor and mineral horizons varied between the three seral age classes following wildfire. Significant differences in mineralizable N, available NO₃⁻-N and available NH₄⁺-N levels with respect to time indicated that available soil N content changes after forest fire. Percent tree and shrub cover was significantly correlated to the amount of available NH₄⁺-N and mineralizable N contents in the forest floor. In the mineral horizons, percent tree cover was significantly correlated to the available NH₄⁺-N, while herb cover was significantly correlated with available NO₃⁻-N. Moss cover was significantly correlated with total N, available NO₃⁻-N and mineralizable N in the forest floor and available NO₃⁻-N in the mineral horizons. We identified several unique species of shrubs and herbs for each seral age class and suggest that plant species are most likely influencing the soil N levels by their contributions to the chemical composition and physical characteristics of the organic matter.     

Biomass and carbon pools of disturbed riparian forests

Quantification of carbon pools as affected by forest age/development can facilitate riparian restoration and increase awareness of the potential for forests to sequester global carbon. Riparian forest biomass and carbon pools were quantified for four riparian forests representing different seral stages in the South Carolina Upper Coastal Plain. Three of the riparian forests were recovering from disturbance (thermal pollution), whereas the fourth represents a mature, relatively undisturbed riparian forest. Above and belowground carbon pools were determined from linear transects established perpendicular to the main stream channels and spanning the width of the riparian area. The objective of this study was to quantify the biomass and carbon pools in severely disturbed, early successional bottomland hardwood riparian forests and to compare these values to those of a less disturbed, mature riparian forest.

Aboveground biomass in all four riparian forests increased during the 2.5-year investigation period. The total carbon pool in these South Carolina Coastal Plain riparian forests increased with forest age/development due to greater tree and soil carbon pools. The mature riparian forest stored approximately four times more carbon than the younger stands. The importance of the herbaceous biomass layer and carbon pool declined relative to total aboveground biomass with increasing forest age. As stands grew older fine root biomass increased, but an inverse relationship existed between percentages of fine root biomass to total biomass. The root carbon pool increased with forest age/development due to a combination of greater fine root biomass and higher root percent carbon.

Aboveground net primary production (NPP) in young riparian forests rapidly approached and exceeded NPP of the more mature riparian forest. As a woody overstory became established (after 8–10 years) annual litterfall rate as a function of NPP was independent of forest age and litterfall amount in the young riparian forests was comparable to mature riparian forests. Biomass in the riparian forest floor and carbon pool declined with increasing riparian forest development. Woody debris in these riparian forests comprised a relatively small carbon pool. An understanding of bottomland hardwood riparian forest carbon pools at different stages of succession allows us to assess how time since disturbance influences these pools, leading to a better understanding of the recovery processes.     

林火对森林演替动态的影响及其应用

森林火灾是危害森林的一个重要的自然因子,但中等强度的林火干扰又是森林生态系统的一个建设因子,林火与森林演替动态的关系密切相关.通过查阅国内外有关林火与森林演替之间相互关系的资料,阐述了林火对森林演替动态的影响,分析了林火与原生演替、次生演替、进展演替和逆行演替的相互关系,列举了林火在维持偏途演替顶极群落和火顶极群落中的作用.     

Long-term influence of fire and harvesting on boreal forest age structure and forest composition in eastern Québec

In boreal forests, historical variations in the area disturbed by natural disturbances or harvesting have rarely been compared. We measured temporal and spatial variations in areas affected by severe fires and clearcutting throughout the 20th century in a 57, 332 km² section of the eastern Canadian boreal forest. We examined the effects of these disturbances on spatio-temporal variations in the abundance of forests >60 years. Natural variability for the abundance of forests >60 years was estimated from simulations of natural disturbance regimes. We also measured compositional and structural differences between three categories of stands originating from relatively recent disturbances (∼50 years; clearcutting, fires, and clearcutting followed by fires), and one category of stands that were undisturbed for at least 200 years. At the regional level, we observed that forests >60 years gradually became scarcer throughout the 20th century due to a gradual expansion of harvested areas, an effect most pronounced in the southern part of the region, where mature and old forest abundance was clearly outside the range of natural variability at the end of the studied period. At the stand level, forest composition and structure differed between stand-origin categories: clearcutting-origin stands contained more balsam fir (Abies balsamea), fire-origin stands more black spruce (Picea mariana), and fire/clearcutting-origin stands more hardwoods (Betula papyrifera and Populus tremuloides). Overall, we estimate that strict forest management targets based on natural disturbance regimes will be difficult to achieve in eastern North-American boreal forests, most notably because contemporary disturbance rates, including both clearcutting and fire, have gradually become higher than the fire rates observed during the preindustrial period.     

森林中的火干扰研究综述

介绍火灾资料分析、火疤木年轮分析等有关火干扰的研究方法.从林火干扰对森林更新、全球气候、森林景观结构维持生物多样性等几方面分析火干扰对森林的影响,认为火干扰的作用具有双重性.高强度火会导致森林生态系统内各种生物死亡,易使林木遭受病虫危害,从而破坏生态系统平衡;而低强度、小面积火的作用有利于改善森林环境,维持森林生态系统平衡,促进森林演替.     

Disturbance history affects lightning fire initiation in the mixedwood boreal forest: Observations and simulations

The emulation of natural disturbances such as fire is a prominent harvest management strategy for ecosystems in Canada's boreal forest region, but the effect of harvesting on subsequent lightning fire occurrence has not been studied systematically in the mixedwood boreal forest. We quantified the relationship between annual patterns of lightning fire initiation, forest composition, lightning, and fire weather conditions over eight years (1994–2001) in a 60,000 km² region of actively harvested mixedwood boreal forest in western Canada. Our analyses illustrated that forest harvesting and burning had opposite effects on subsequent fire initiation, so harvest was not a surrogate for fire. Fire initiation increased in landscapes with more area harvested and decreased with area recently burned. Our data suggested that increased fire initiation was most pronounced in harvested stands up to a decade old, and there was some evidence that the effect might last as long as 30 years. We then used a dynamic fire-succession simulation model to quantify the long-term effects of these fuel-based relationships using two metrics. As expected, the first metric demonstrated that the number of years between disturbances was significantly less in stands that were harvested and then burned, than those that were burned and then burned again. However, the more revealing component of the simulations was an illustration that despite the strong, positive relationship between harvested areas and fire initiation, the area affected over the long-term by a reduced disturbance interval was relatively small. Accordingly, our study shows that spatiotemporal regulation of lightning fire initiation through harvesting activity results in a systematic accelerated frequency of disturbance that is novel to the mixedwood boreal system, but the area affected by these events amount to local peculiarities rather than broad-scaled regularities.     

不同干扰类型下的白桦次生林土壤理化性质研究

研究内蒙古大兴安岭地区不同干扰类型下白桦次生林土壤理化性质特征,在内蒙古根河市选取3种干扰类型的白桦次生林样地的不同土层土壤作为研究对象,对土壤理化性质进行测量分析,探究不同干扰类型对土壤理化性质的影响。研究结果表明,白桦次生林在不同干扰方式下土壤理化性质存在显著差异。其中人工条带改造后的白桦次生林土壤pH值增大,火烧后白桦次生林容重增大,土壤有机碳、全氮、全磷、全钾、速效磷、速效钾含量在人工条带改造后显著增高,采伐干扰样地白桦次生林样地土壤含水量、速效氮含量最高。不同干扰类型的白桦次生林样地土壤理化性质间相关性不同。人工条带改造干扰样地白桦次生林土壤综合指数较高,火干扰样地白桦次生林土壤综合指数较低。     

Succession of mosses,liverworts and ferns on coarse woody debris,in relation to forest age and log decay in Tasmanian wet eucalypt forest

In managed forest landscapes, understanding successional processes is critical to management for sustainable biodiversity. Coarse woody debris is a key substrate for forest biodiversity, particularly because it undergoes complex succession reflecting the effects of changes in both forest structure and substrate characteristics. The present study used a chronosequence approach to investigate succession of mosses, liverworts and ferns on coarse woody debris following clearfell, burn, sow native forest silviculture in wet eucalypt forest in Tasmania, focussing on discriminating between the effects of forest age and log decay. It also compared successional processes following wildfire with those following clearfell, burn, sow silviculture. Forest regenerating after the latter form of regeneration showed clear ecological succession up to 43 years (the limit of available sites), characterised by increasing diversity and cover, and clearly delineated specialisation among species with regard to successional stage. Analyses of subsets of the full data-set indicated that the effects of forest age dominated this succession, with minimal effects of substrate change independent of forest age. Analysis of within forest microenvironments were consistent with the inference that microenvironmental changes related to forest age drive major successional changes in these forests. Comparative analysis indicated similarity between successional states in post-wildfire and post- clearfell, burn, sow regeneration after 43 years for logs of the same decay stages, and continuing succession on post-wildfire sites to at least 110 years. Overall, these data suggest that management to sustain fern and bryophyte diversity should ensure that areas of forest beyond 110 years are represented in the landscape at appropriate spatial scales.