A spatial simulation model for forest succession in the Upper Mississippi River floodplain

A Markov-chain transition model (FORSUM) and Monte Carlo simulations were used to simulate the succession patterns and predict a long-term impact of flood on the forest structure and growth in the floodplain of the Upper Mississippi River and Illinois River. Model variables, probabilities, functions, and parameters were derived from the analysis of two comprehensive field surveys conducted in this floodplain. This modeling approach describes the establishment, growth, competition, and death of individual trees for modeled species on a 10,000-ha landscape with spatial resolution of 1 ha. The succession characteristics of each Monte Carlo simulation are summed up to describe forest development and dynamics on a landscape level. FORSUM simulated the impacts of flood intensity and frequency on species composition and dynamics in the Upper Mississippi River floodplain ecosystem. The model provides a useful tool for testing hypotheses about forest succession and enables ecologists and managers to evaluate the impacts of flood disturbances and ecosystem restoration on forest succession. The simulation results suggest that the Markov-chain Monte Carlo method is an efficient tool to help organize the existing data and knowledge of forest succession into a system of quantitative predictions for the Upper Mississippi River floodplain ecosystem.     

林窗干扰与森林群落演替

林窗干扰是影响森林群落演替的一个重要因素。该文从林窗干扰和森林群落演替理论及林窗对森林群落微环境、植物入侵和定居、群落结构、群落演替的影响等方面简要介绍了当前国内外有关林窗干扰与森林群落演替的研究现状和研究前景。     

Non-native grass invasion suppresses forest succession

Multiple factors can affect the process of forest succession including seed dispersal patterns, seedling survival, and environmental heterogeneity. A relatively understudied factor affecting the process of succession is invasions by non-native plants. Invasions can increase competition, alter abiotic conditions, and provide refuge for consumers. Functional traits of trees such as seed size and life history stage may mediate the effects of invasions on succession. We tested the effects of the forest invader Microstegium vimineum on planted and naturally regenerating trees in a multi-year field experiment. We established plots containing nine species of small- and large-seeded tree species planted as seeds or saplings, and experimentally added Microstegium to half of all plots. Over 3 years, Microstegium invasion had an overall negative effect on small-seeded species driven primarily by the effect on sweetgum, the most abundant small-seeded species, but did not affect large-seeded species such as hickory and oak species, which have more stored seed resources. Natural regeneration was over 400% greater in control than invaded plots for box elder, red maple, and spicebush, and box elder seedlings were 58% smaller in invaded plots. In contrast to the effects on tree seedlings, invasion did not affect tree sapling survival or growth. Microstegium may be directly reducing tree regeneration through competition. Invaded plots had greater overall herbaceous biomass in 2006 and 2008 and reduced light availability late in the growing season. Indirect effects may also be important. Invaded plots had 120% more thatch biomass, a physical barrier to seedling establishment, and significantly greater vole damage to tree saplings during 2006 and 2007. Our results show that two tree functional traits, seed size and life history stage, determined the effects of Microstegium on tree regeneration. Suppression of tree regeneration by Microstegium invasions may slow the rate of forest succession and alter tree species composition.     

气候变化对森林演替的影响

森林演替是森林生态动力源驱动下森林再生的生态学过程,自20世纪初建立群落演替理论以来,演替研究成为生态学研究中的热点．客观准确地认识森林演替规律,研究森林演替动力学机理及其模型,是科学管理森林生态系统的需要;对于天然林保护工程与森林植被的恢复重建,具有重要的理论与实际意义．干扰是森林循环的驱动力,导致森林生态系统时空异质性,是更新格局和生态学过程的主要影响因素．它可改变资源的有效性,干扰导致的林隙是森林循环的起点．回顾了目前演替研究的几种方法,即马尔科夫模型、林窗模型(GAP)、陆地生物圈模型(BIOME)和非线性演替模式．介绍了气候变化对森林演替的影响;并在已有成果的基础上,提出了目前研究存在的问题及未来的发展方向．     

A stoichiometric model of early plant primary succession

The relative importance of plant facilitation and competition during primary succession depends on the development of ecosystem nutrient pools, yet the interaction of these processes remains poorly understood. To explore how these mechanisms interact to drive successional dynamics, we devised a stoichiometric ecosystem-level model that considers the role of nitrogen and phosphorus limitation in plant primary succession. We applied this model to the primary plant community on Mount St. Helens, Washington State, to check the validity of the proposed mechanisms. Our results show that the plant community is colimited by nitrogen and phosphorus, and they confirm previous suggestions that the presence of a nitrogen-fixing legume, Lupinus lepidus, can enhance community biomass. In addition, the observed nutrient supply rates may promote alternative successional trajectories that depend on the initial plant abundances, which may explain the observed heterogeneity in community development. The model further indicates the importance of mineralization rates and other ecosystem parameters to successional rates. We conclude that a model framework based on ecological stoichiometry allows integration of key biotic processes that interact nonlinearly with biogeochemical aspects of succession. Extension of this approach will improve the understanding of the process of primary succession and its application to ecosystem rehabilitation.     

Functional diversity changes during tropical forest succession

Functional diversity (FD) ‘those components of biodiversity that influence how an ecosystem operates or functions’ is a promising tool to assess the effect of biodiversity loss on ecosystem functioning. FD has received ample theoretical attention, but empirical studies are limited. We evaluate changes in species richness and FD during tropical secondary forest succession after shifting cultivation in Mexico. We also test whether species richness is a good predictor of FD. FD was calculated based on a combination of nine functional traits, and based on two individual traits important for primary production (specific leaf area) and carbon sequestration (wood density). Stand basal area was a good predictor of successional changes in diversity and FD, in contrast to fallow age. Incidence-based FD indices increased logarithmically with stand basal area, but FD weighted by species’ importance values lacked pattern with succession. Species richness and diversity are strong predictors of FD when all traits were considered; linear relationships indicate that all species are equally functionally complementary, suggesting there is little functional redundancy. In contrast, when FD was calculated for individual traits and weighted for abundances, species richness may underestimate FD.Selection of functional trait(s) critically determines FD, with large consequences for studies relating biodiversity to ecosystem functioning. Careful consideration of the traits required to capture the ecosystem process of interest is thus essential.     

Causes of arrested succession in coastal dune forest

Dispersal or recruitment limitation may arrest succession after disturbance. In north-eastern South Africa the Acacia karroo successional pathway is used to facilitate coastal forest recovery after strip-mining. However, although A. karroo establishes naturally, it forms monospecific stands, arresting forest succession for decades. This casts doubt on the efficacy of this restoration pathway. We investigated the causes of arrested succession. The seed and seedling banks of A. karroo stands and of forest at Cape Vidal, and three A. karroo stands (7–27 years old) on rehabilitated strip-mined dunes at nearby Richards Bay were examined. The establishment and growth of seedlings at Cape Vidal were also considered. The seed bank was larger and more diverse in forest, but the seedling bank was larger in Acacia stands. At Richards Bay, the size of the seed bank increased and the seedling bank decreased with Acacia stand age. Excluding mammalian herbivores in Acacia stands at Cape Vidal resulted in greater species richness and survival of naturally established seedlings, as well as two experimentally planted species. Neither seed dispersal nor seedling establishment limited recruitment of tree species in Acacia stands. Herbivory arrested forest succession by causing the differential mortality of seedlings. In contrast, at Richards Bay where there were few mammalian herbivores, the advanced regeneration in A. karroo stands converged on the diversity of nearby forests 29 years after restoration. Controlling herbivore access and seeding Acacia stands with forest species are site-specific options for preventing arrested succession when using the A. karroo successional pathway.     

Ordination of simulated complex forest succession: A new test of ordination methods

A model of a 1/12th ha forest stand, FORET, generated 10 000 years of simulated species succession. Approximately the first third of these results were analyzed by principal component analysis as if they were collected field data to give the trajectory of the community particle in a collapsed species space. The ordination axis orientation was performed on a dispersion matrix and correlation matrix between species. In both cases, however, the eigen vectors were applied to the data matrix which had not been transformed to unit species variance. This facilitated comparison of species dispersion and correlation structure; it emerged they were very different. Correlation structure gave large weights to understory species while dispersion emphasized the dominant overstory species. This implies a decomposition of simulated stand behavior into overstory and understory, even though such decomposition was not formally built into the model. This decomposition would seem to pertain to real vegetation. Principal component analysis was able to express insightful differences between data structure with and without the unit variance transformation implicit in the correlation matrix. This flexibility of the ordination method proved valuable in uncovering unsuspected ordering principles in the model. Complex simulated data allow the ordination technique to demonstrate its capacity to generate new hypotheses, which hypotheses can then be simply validated by a return to the structure of the model but with the hindsight of the analysis. The generation of new hypotheses is not possible if the simulation is of a simple coenocline; on the other hand, ordination of test field data does not allow the simple validation of new hypotheses, for in the field there is not a defined algorithm to which the researcher can return.     

Long-term succession in a Danish temperate deciduous forest

Forest successional trajectories covering the last 2000 yr from a mixed deciduous forest in Denmark show a gradual shift in dominance from Tilia cordata to Fagus sylvatica and a recent increase in total forest basal area since direct management ceased in 1948. The successions are reconstructed by combining a fifty-year record of direct tree observations with local pollen diagrams from Draved Forest, Denmark. Five of the seven successions record a heathland phase of Viking Age dating from 830 AD. The anthropogenic influence is considerable throughout the period of study even though Draved contains some of the most pristine forest stands in Denmark. Anthropogenic influence including felling masks the underlying natural dynamics, with the least disturbed sites showing the smallest compositional change. Some effects of former management, such as loss of Tilia cordata dominance, are irreversible. Artificial disturbance, particularly drainage, has accelerated and amplified the shift towards Fagus dominance that would have occurred on a smaller scale and at a slower rate in the absence of human intervention.     

Seed dynamics during forest succession in Costa Rica

Soil seed banks and current seed inputs each play a role in tropical succession. We compared the abundance and floristic composition of seeds from these two sources at a Costa Rican site by germinating seeds from the soil, measuring seed inputs for 3 yr, and monitoring the earliest colonists in a forest clearing.There were an estimated 6800 viable seeds/m² in the soil of 3.3-yr-old vegetation, 9500 seeds/m² in 11-yr-old vegetation, and 7000 seeds/m² in a 75-yr-old forest. An estimated 10100 seeds/m² fell on the soil surface of the young successional vegetation during 3 yr and 3700 seeds/m² fell during that same time in the forest.Locally produced seeds accounted for about 75% of the seed input to the soil surface early in succession. Seeds dispersed out of young successional vegetation increased the quantity and species richness of the seed input and storage in an adjacent forest. Much of the species richness of the young successional vegetation resulted from seeds dispersed there from other communities by animals.Deforestation stimulated germination of most seeds in the surface soil of the old forest, including seeds of the dominant canopy tree. The recruitment of seedlings from the soil seed bank numerically overwhelmed that from post-disturbance seed rain and sprouts.We evaluated patterns of soil seed storage during succession and predicted the ability of vegetation of differing ages to respond to disturbance. Immediately after disturbance the number of seeds in the soil plummeted due to mortality, low inputs, and germination. As the vegetation regrew, the soil seed bank increased to a peak after 4 to 7 yr, then gradually decreased to its pre-disturbance size. High-frequency pulses of disturbance should result in reduced species richness, dominance by species with long-lived seeds, and fast recovery by seedling recruitment from the soil seed bank.Journal series number 6459 from the Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida 32611, USA.Reprint requests to J. J. E. at Florida.     

Landslide early succession in a neotropical dry forest

Early succession on large landslides in highly humanized areas that have a tropical dry climate is not well studied. This study documented vegetation recovery during the first 4 years after disturbance at a landslide on Casita Volcano, Nicaragua. We aimed to determine the main pathways and causes of change in community features, such as richness, biovolume, and species composition and verify the role played by environmental heterogeneity. Data consisting on number, covers and mean height of woody species and several abiotic factors related to fertility and stability of substrates were obtained from permanent plots in previously defined zones. Pathways of early succession were highly contingent on abiotic heterogeneity and landscape context and were mainly controlled by abiotic factors associated with fertility of substrates, and incidence of human disturbances. Those results might form the basis of a model of early succession on landslides located in densely populated areas within tropical dry ecosystems. Our results suggest that, rather than focusing research on large-scale disturbances, the study of succession in landslides of the type that occurred on Casita Volcano must point towards the response of ecosystems to a much more complex disturbance regime, in which human-induced disturbances play a major role.     

Phylogenetic structure of angiosperm communities during tropical forest succession

The phylogenetic structure of ecological communities can shed light on assembly processes, but the focus of phylogenetic structure research thus far has been on mature ecosystems. Here, I present the first investigation of phylogenetic community structure during succession. In a replicated chronosequence of 30 sites in northeastern Costa Rica, I found strong phylogenetic overdispersion at multiple scales: species present at local sites were a non-random assemblage, more distantly related than chance would predict. Phylogenetic overdispersion was evident when comparing the species present at each site with the regional species pool, the species pool found in each age category to the regional pool or the species present at each site to the pool of species found in sites of that age category. Comparing stem size classes within each age category, I found that during early succession, phylogenetic overdispersion is strongest in small stems. Overdispersion strengthens and spreads into larger size classes as succession proceeds, corroborating an existing model of forest succession. This study is the first evidence that succession leaves a distinct signature in the phylogenetic structure of communities.     

子午岭次生林植被演替过程的土壤抗冲性

在水土流失极为严重的黄土高原地区,土壤抗冲性决定着土壤的可蚀性.于2004年5月在黄土高原惟一的次生林区——子午岭林区,通过原状土冲刷实验对不同植被演替阶段下的土壤抗冲性进行了研究.结果表明： （1）随植被的正向演替,表层土壤（0～15cm）的抗冲性明显增大,但亚表层（15～30cm）和底层（30～50cm）土壤抗冲性则没有太大的变化;（2）植物根系能显著的增强土壤抗冲性,土壤抗冲系数与单位土体根系表面积具有极显著的（p＜0.001）线性相关关系;（3）土壤抗冲系数随土壤中水稳性团聚体含量和微生物量的增加而增大,且其相关关系极显著（p＜0.001）.综合根系（x1）、水稳性团聚体（x2）以及微生物（x3）对土壤抗冲性的影响,建立黄土高原地区土壤抗冲性方程： y=-4.89＋1.27x1＋0.079x2＋1.94E-3x3 （R^2=0.914 p＜0.001）.     

The role of frugivorous bats in tropical forest succession

Discussion of successional change has traditionally focused on plants. The role of animals in producing and responding to successional change has received far less attention. Dispersal of plant propagules by animals is a fundamental part of successional change in the tropics. Here we review the role played by frugivorous bats in successional change in tropical forests. We explore the similarities and differences of this ecological service provided by New and Old World seed-dispersing bats and conclude with a discussion of their current economic and conservation implications. Our review suggests that frugivorous New World phyllostomid bats play a more important role in early plant succession than their Old World pteropodid counterparts. We propose that phyllostomid bats have shared a long evolutionary history with small-seeded early successional shrubs and treelets while pteropodid bats are principally dispersers of the seeds of later successional canopy fruits. When species of figs (Ficus) are involved in the early stages of primary succession (e.g. in the river meander system in Amazonia and on Krakatau, Indonesia), both groups of bats are important contributors of propagules. Because they disperse and sometimes pollinate canopy trees, pteropodid bats have a considerable impact on the economic value of Old World tropical forests; phyllostomid bats appear to make a more modest direct contribution to the economic value of New World tropical forests. Nonetheless, because they critically influence forest regeneration, phyllostomid bats make an important indirect contribution to the economic value of these forests. Overall, fruit-eating bats play important roles in forest regeneration throughout the tropics, making their conservation highly desirable.     

A computer model of succession and fire response of the high-altitude Eucalyptus forest of the Brindabella Range,Australian Capital Territory

The BRIND model, a computer model of the high altitude forests in the Brindabella Range near Canberra (Australian Capital Territory), is documented and the results of a series of tests on the model are provided. The BRIND model simulates a 1/12 ha forest stand by computing the growth of each individual tree in the stand. It considers establishment and death of trees on a tree-by-tree basis using stochastic functions. The model also simulates the effects of prescribed fire and wildfire on the forests. The model presently is restricted to southeasterly facing slopes (moist, sheltered situations) above 850 m in altitude. The BRIND model is tested in four ways: (1) A single example simulation (for 500years) is inspected for agreement with stand dynamics in wet sclerophyll forests. (2) By varying wildfire frequency, the model is used to develop a succession diagram for forests in the alpine ash (E. delegatensis) zone of the Brindabella Range. This diagram is considered in terms of the successional patterns described for this ecological zone. (3) By subjecting the model to different climatic conditions and wildfire frequencies, a simulated altitude zonation is developed. This simulated pattern of forest types is compared with the extant forest types in the Brindabella Range. (4) The model is tested on its ability to duplicate basal area, stocking density, and average diameters for different age stands found in an independent data set. The model was found to simulate patterns of vegetation that resemble those of the forests of the Brindabella Range in both space and time. The successional pattern was found to be complex and to differ from classic theories of succession originating with Clements. Potential model applications are discussed     

广西木荷林的分类和演替

木荷（ＳｃｈｉｍａｓｕｐｅｒｂａＧａｒｄｎ．ｅｔＣｈａｍｐ．）林是原来的常绿阔叶林遭受破坏以后,在保护较好的情况下,从演替系列群落顺向演替恢复为常绿阔叶林的一个早期类型。本文对广西木荷林进行分类,划分和论述７个群丛的种类组成特点和演替方向,可供进一步研究和规划退化生态系统的恢复和重建参考     

Incorporating environmental change over succession in an integral projection model of population dynamics of a forest herb

Despite seemingly obvious effects of environmental drivers, mechanisms behind long‐term changes in plant population sizes over time are often poorly known. We investigated how soil potassium concentration and seed predation are likely to change over time as a result of succession from deciduous forest to spruce forest, and how this affects population trajectories of Actaea spicata. Observations and addition experiments showed that high soil potassium concentration increased individual growth rates. Among‐site comparisons showed that soil potassium concentration was lower where proportion spruce was higher. Incorporation of a gradual increase in spruce over time in an integral projection model where individual growth depended on potassium suggested a net decrease in A. spicata population sizes over forest succession. This result suggests that small changes in factors with small effects on individual performance can influence patterns of species occupancy along successional gradients. We incorporated also density independent and density dependent effects of pre‐dispersal seed predation over succession into the same model. Seed predation influenced the tree composition at which A. spicata population growth was positive. However, significant effects of A. spicata population size on seed predation intensity did not translate into important feedback effects on population growth trajectories over succession. Our results illustrate how demographic models can be used to gain understanding of the mechanisms behind effects of environmental change on species abundances and distributions by the simultaneous inclusion of changing abiotic and biotic factors.     

A simple mathematical model of the secondary succession of shrubs

A simple simulation model of the secondary shrub succession has been elaborated on the grounds of primary field data from abandoned fields of different ages in the area of Bohemian Karst. The model describes vegetational dynamics using adaptedVolterra-Lotka equations for competing species. Carrying capacities and growth rates are expressed as a function of the depth of the soil.     

Quantifying ecological memory during forest succession: A case study from lower subtropical forest ecosystems in South China

The concept of ecological memory provides a new perspective for research on forest succession by including historical factors and the initial state of ecological processes. However, there are still significant gaps between the concept and its application. We selected nine proxy indicators (plant species, soil seed banks, soil microbes, soil animals, birds, soil age, soil pollen, soil mineral distribution, and light environment) and developed a method to quantify ecological memory and succession in a subtropical forest succession in South China. Taking the climax-monsoon evergreen broad-leaved forest as the reference ecosystem, we found that ecological memory increased nonlinearly and accumulated following a specific assembly rule during succession. Memory concerning major soil microbes and soil animals, which improve the soil substrate, mainly accumulated from the initial to the early successional stage. Memory concerning the number of bird species and the availability of light, which ensure a source of regenerative seeds and the survival of understory seedlings, mainly accumulated from the early to middle successional stages. Memory concerning vegetation and soil seed banks mainly accumulated late in succession, guaranteeing that the ecosystem would reach the regional climax stage. Prospective memory was greater than retrospective memory in every successional stage except the late stage, which indicated that all stages but the late stage were undergoing progressive succession. Our study demonstrates that the concept of ecological memory and the proposed evaluation framework are useful for guiding research on succession and restoration, and especially for assessing how “far” a restored ecosystem is from a reference ecosystem or how far a restored ecosystem has deviated from its natural succession trajectory.     

森林演替在南亚热带森林生态系统碳吸存中的作用

研究了鼎湖山南亚热带森林同一演替系列中3个不同演替阶段(马尾松针叶林、马尾松荷木混交林和季风常绿阔叶林)生态系统碳贮量和分配格局特征,并探讨了该地区森林演替过程中生态系统碳吸存潜力和速度。结果表明:(1)针叶林各组分碳素含量高于阔叶林对应组分的碳素含量(后者是前者的72.0%～94.5%)。两个森林植物碳素含量,不同层次比较,均为乔木层>灌木层>草本层,不同器官比较,以根或干最高。(2)乔木层生物量随森林演替进展而增加。针叶林、混交林和阔叶林乔木层生物量分别为:143.5t/hm2、270.1t/hm2和407.8t/hm2,其中大部分由干和皮组成(各器官占乔木层生物量的比例平均为:叶2.8%、枝19.3%、干和皮混合57.0%、根20.9%)。林下层生物量为4.23～14.10t/hm2,是乔木层的1.0%～9.8%,随森林演替进展而减少。(3)土壤容重随深度增加而增加,但随森林演替进展而减少。与土壤容重相反,土壤有机碳含量随深度增加而明显减少,但随森林演替进展而增加。(4)3种类型森林生态系统碳总贮量分别为135.8t/hm2、215.1t/hm2和259.7t/hm2。生态系统碳贮量在各组分的格局十分相似,植被、土壤和凋落物层所占比例均分别约为67.6%、30.2%和2.2%。与其它地带森林比较,鼎湖山保护区森林植被与土壤碳贮量之比和表层(0～20cm)的土壤碳占整个