Successional theories

Succession is a fundamental concept in ecology because it indicates how species populations, communities, and ecosystems change over time on new substrate or after a disturbance. A mechanistic understanding of succession is needed to predict how ecosystems will respond to land-use change and to design effective ecosystem restoration strategies. Yet, despite a century of conceptual advances a comprehensive successional theory is lacking. Here we provide an overview of 19 successional theories (‘models’) and their key points, group them based on conceptual similarity, explain conceptual development in successional ideas and provide suggestions how to move forward. Four groups of models can be recognised. The first group (patch & plants) focuses on plants at the patch level and consists of three subgroups that originated in the early 20th century. One subgroup focuses on the processes (dispersal, establishment, and performance) that operate sequentially during succession. Another subgroup emphasises individualistic species responses during succession, and how this is driven by species traits. A last subgroup focuses on how vegetation structure and underlying demographic processes change during succession. A second group of models (ecosystems) provides a more holistic view of succession by considering the ecosystem, its biota, interactions, diversity, and ecosystem structure and processes. The third group (landscape) considers a larger spatial scale and includes the effect of the surrounding landscape matrix on succession as the distance to neighbouring vegetation patches determines the potential for seed dispersal, and the quality of the neighbouring patches determines the abundance and composition of seed sources and biotic dispersal vectors. A fourth group (socio-ecological systems) includes the human component by focusing on socio-ecological systems where management practices have long-lasting legacies on successional pathways and where regrowing vegetations deliver a range of ecosystem services to local and global stakeholders. The four groups of models differ in spatial scale (patch, landscape) or organisational level (plant species, ecosystem, socio-ecological system), increase in scale and scope, and reflect the increasingly broader perspective on succession over time. They coincide approximately with four periods that reflect the prevailing view of succession of that time, although all views still coexist. The four successional views are: succession of plants (from 1910 onwards) where succession was seen through the lens of species replacement; succession of communities and ecosystems (from 1965 onwards) when there was a more holistic view of succession; succession in landscapes (from 2000 onwards) when it was realised that the structure and composition of landscapes strongly impact successional pathways, and increased remote-sensing technology allowed for a better quantification of the landscape context; and succession with people (from 2015 onwards) when it was realised that people and societal drivers have strong effects on successional pathways, that ecosystem processes and services are important for human well-being, and that restoration is most successful when it is done by and for local people. Our review suggests that the hierarchical successional framework of Pickett is the best starting point to move forward as this framework already includes several factors, and because it is flexible, enabling application to different systems. The framework focuses mainly on species replacement and could be improved by focusing on succession occurring at different hierarchical scales (population, community, ecosystem, socio-ecological system), and by integrating it with more recent developments and other successional models: by considering different spatial scales (landscape, region), temporal scales (ecosystem processes occurring over centuries, and evolution), and by taking the effects of the surrounding landscape (landscape integrity and composition, the disperser community) and societal factors (previous and current land-use intensity) into account. Such a new, comprehensive framework could be tested using a combination of empirical research, experiments, process-based modelling and novel tools. Applying the framework to seres across broadscale environmental and disturbance gradients allows a better insight into what successional processes matter and under what conditions.     

How Much Ecology Do We Need to Know to Restore Mediterranean Ecosystems?

Despite important advances in ecological knowledge of Mediterranean‐type ecosystems, advances in restoration ecology have not seen a parallel increase in these systems. Although some concepts such as positive plant–plant interaction (facilitation) have received attention in the restoration ecology community, others such as phenotypic plasticity have not. Some concepts (e.g., environmental heterogeneity) are mature enough for a wide use in restoration, whereas available knowledge on others (e.g., facilitation, plasticity) is less conclusive. However, the scientific knowledge is in general enough to significantly improve the guidelines for restoration of Mediterranean ecosystems. Our review suggests that (1) the extent of facilitation in dry ecosystems is partially understood, with supporting, but somewhat contradictory empirical evidence for its potential use in restoration; (2) the influence of habitat heterogeneity on plant performance and plasticity is only beginning to be understood, with a strong bias toward patterns of structural heterogeneity and negligible information on functional heterogeneity; and (3) sound evaluations of phenotypic plasticity might be useful to increase the success of restoration practices in patchy Mediterranean environments. Future global change scenarios involving temperature rise, reduced precipitation, increased frequency of extreme climatic events, and important land use changes and fragmentation must be particularly considered when restoring Mediterranean ecosystems. Further research on how to incorporate results on facilitation, environmental heterogeneity, and plasticity within a global change framework is clearly needed.     

Transfer seeds,hay, or soil blocks? The importance of the completeness of biological inputs to address dispersal and establishment limitations during the restoration of plant assemblages in floodplain grasslands

This study investigated different techniques of grassland restoration to overcome dispersal or establishment limitation, which are key processes influencing early-successional plant community assembly. A fully randomized in situ experiment was set up in a former arable land in a floodplain along the Garonne river (south-western France) to test for the effect of (1) the type and completeness of the biological input (any biological material—seed, hay, and soil—transferred to the site under restoration), (2) soil disturbance by deep tillage, and (3) their interaction on plant community dynamics for 5 years. All inputs influenced the plant community structure and composition. The effects depended on the type of the input but not necessarily in link with its completeness. High density and diversity seed mixture led to high levels of richness and relative abundance of target species like soil blocks, the input considered as the most complete. During the first year, hay transfer mostly influenced community assembly through negative litter effects. Delayed germination of several species contributed to buffer these early effects. This study supports the importance of dispersal limitation during early succession in degraded grassland ecosystems whereas soil disturbance had only subtle effects on the seed bank and standing plant community, indicating that competition and establishment limitation were of secondary importance in our study system. Our results suggest that even low-frequency immigration events (e.g. when a few seeds are transferred within hay) can be of great importance providing that a sufficiently long time period is considered for grassland restoration.     

Protocols for Restoration Based on Recruitment Dynamics, Community Structure, and Ecosystem Function: Perspectives from South African Fynbos

South African fynbos ecosystems are under threat from alien plant invasions and transformations to alternative land uses. If large-scale habitat loss and species extinction are to be halted, restoration actions are urgently required. We postulate that by adopting an approach in which an understanding of community and ecosystem dynamics is applied to restoration practices, protocols can be developed which will lead to more efficient restoration. This understanding is based on a review of the relevant ecological literature, focusing on recruitment dynamics, community structure, and ecosystem function, which are particularly relevant to restoration. We develop a conceptual framework for restoration and apply our protocols to a case study area on the Cape Peninsula. Before ecological restoration can begin, the cause of transformation must first be removed or ameliorated. The next step is to ensure that the important ecological processes are functioning. We contend that a fully functioning community requires a good balance of the major growth-form, regeneration, and nutrient acquisition guilds. Fire is the natural disturbance event initiating recruitment in fynbos. It is, therefore, essential to either burn a site or provide fire-related germination cues in order to stimulate recruitment. Where guilds are under-represented, corrective reintroductions will further improve the long-term resilience of the restored community. Many taxa have persistent soil-stored seed banks, so it is important to conserve topsoil and optimize use of this local species pool. Seed dispersal distances are generally very short, and in highly transformed sites it will be necessary to reintroduce seed of the major guilds in order to restore community structure and functioning. Post-fire succession in fynbos begins with the full complement of species; species gradually die out from the vegetation according to their respective life spans. In order to stimulate germination and promote successful establishment, it is important to sow seed after fire or site clearing in late summer or autumn. Introducing seed or plants at a later stage in the succession is very likely to fail. Because of the localized distributions of many taxa, extreme care must be taken when selecting species for reintroduction if local gene pools are to be conserved.     

Generation of Spatial Patterns in Boreal Forest Landscapes

Boreal forests are composed of a few plant species with contrasting traits with respect to ecosystem functioning and spatial patterning. Early successional deciduous species, such as birch and aspen, disperse seeds widely, do not tolerate low light and nitrogen availabilities, have rapidly decaying litter, and are highly preferred by herbivores. These later succeed to conifers, such as spruce and fir, which disperse seeds locally, tolerate low light levels and low nitrogen availability, have litter that decays slowly, and are unpalatable to most mammalian herbivores. Although there are also early successional conifers, such as jack pine and Scots pine, the aspen-birch-spruce-fir successional sequence is the most common over much of North America, and (without fir) in Fennoscandia and Siberia. The course of succession in these forests is controlled partly by seed dispersal and selective foraging by mammalian herbivores. Both of these processes are spatially dynamic, but little is known about how their spatial dynamics may affect ecosystem processes, such as nitrogen cycling or productivity. We present spatially explicit models that demonstrate the following: (a) Spatially explicit seed dispersal results in more clumped distribution of tree species and persistence of greater paper birch biomass than uniform seed rain across the landscape. Such results are consistent with current spatially explicit population models of dispersal and coexistence. (b) With localized seed dispersal, the concentrations of available soil nitrogen are distributed in larger patches with sharp transitions from low to high nitrogen availability near patch edges. In contrast, with a uniform seed rain, the distribution of soil nitrogen availability was more uniform and “hotspots” were more localized. Thus, the spatial pattern of an ecosystem process (nitrogen cycling) is determined by seed dispersal and competition for light among competing populations. (c) A dispersing herbivore, such as moose, that selectively forages on early successional deciduous species with high quality litter, such as aspen or birch, and discriminates against late successional conifers, such as spruce or fir, imposes higher-order repeated patterns of plant species and biomass distribution on the landscape. Thus, seed dispersal and herbivore foraging correlate properties in adjacent patches but in different ways, and different spatial patterns emerge. Other processes, such as insect outbreaks, fire, and water flow, also may correlate properties between adjacent patches and result in additional patterns. Received 8 February 1999; accepted 28 May 1999.     

Effects of Natural and Human-Assisted Regeneration on Landscape Dynamics in a Korean Pine Forest in Northeast China

Improper forest harvesting can potentially degrade forest ecosystem functions and services. Human-assisted regeneration (e.g., planting) is often used to increase the rate of forest recovery and thereby reduce regeneration failure. Seed dispersal is a fundamental ecological process that can also influence spatio-temporal patterns of forest regeneration. In this study, we investigated the relative contribution of planting and seed dispersal on forest regeneration at landscape scales. Because such influences can be further complicated by timber harvest intensity and seed availability within and around harvested area, we also evaluated the effects of those factors on forest landscape dynamics. We used the forest landscape model LANDIS to simulate the dynamics of Korean pine-broadleaf mixed forests in Northeast China. We considered three factors: timber harvest intensity (3 levels), seed dispersal and whether or not planting was used. The results showed that planting was more important in maintaining the abundance of Korean pine (Pinus koraiensis), a climax keystone species in this region, under the high-intensity harvesting option during early succession. In contrast, seed dispersal was more important during late succession. Korean pine can be successfully regenerated through seed dispersal under low and medium harvest intensities. Our results also indicated that effective natural regeneration will require protecting seed-production trees (seed rain). This study results provide a basis for more effectively managing Chinese temperate forests and possibly other similar ecosystems.     

Succession of Ephemeral Secondary Forests and Their Limited Role for the Conservation of Floristic Diversity in a Human-Modified Tropical Landscape

Both local- and landscape-scale processes drive succession of secondary forests in human-modified tropical landscapes. Nonetheless, until recently successional changes in composition and diversity have been predominantly studied at the patch level. Here, we used a unique dataset with 45 randomly selected sites across a mixed-use tropical landscape in central Panama to study forest succession simultaneously on local and landscape scales and across both life stages (seedling, sapling, juvenile and adult trees) and life forms (shrubs, trees, lianas, and palms). To understand the potential of these secondary forests to conserve tree species diversity, we also evaluated the diversity of species that can persist as viable metapopulations in a dynamic patchwork of short-lived successional forests, using different assumptions about the average relative size at reproductive maturity. We found a deterministic shift in the diversity and composition of the local plant communities as well as the metacommunity, driven by variation in the rate at which species recruited into and disappeared from the secondary forests across the landscape. Our results indicate that dispersal limitation and the successional niche operate simultaneously and shape successional dynamics of the metacommunity of these early secondary forests. A high diversity of plant species across the metacommunity of early secondary forests shows a potential for restoration of diverse forests through natural succession, when trees and fragments of older forests are maintained in the agricultural matrix and land is abandoned or set aside for a long period of time. On the other hand, during the first 32 years the number of species with mature-sized individuals was a relatively small and strongly biased sub-sample of the total species pool. This implies that ephemeral secondary forests have a limited role in the long-term conservation of tree species diversity in human-modified tropical landscapes.     

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.     

恢复及演替过程中的土壤生态学考虑

 人类社会的日益扩张,导致人类加速占据地球表面景观,并胁迫地球上生态系统提供不断增长的资源需求和废物吸收能力。所以保护尚未“开放”的自然生态系统及恢复退化的生态系统成为人类长期生存的重要保证。该文着重讨论了恢复过程中的土壤生态学问题。土壤是所有陆地生态系统的结构与功能基础。土壤微生物与动物的种群变化,土壤有机质的积累,及主要元素地球化学循环的改变是恢复生态的重要环节。生态恢复与演替有许多共性,所以演替理论对于认识生态系统恢复中的结构与功能变化有着很大帮助。与自然演替不同的是,人的积极参与在生态恢复中占有中心位置。从最初样地的确立与物种的选择,到后续的灌溉与施肥管理,人的选择影响着土壤的演化,生态系统的发展方向,和最终恢复生态的结果。为保障恢复生态系统的可持续性,短期的工作目标,如提供养分促进植物生长,务必与长期的工作目标,如土壤的恢复相结合。植物与土壤的相互反馈是生态恢复成功的重要标志。成功的生态恢复不仅是对现有生态学理论的“试金检验”,也是推动生态学学科发展的重要原动力。     

Differences in CO2 dynamics between successional mire plant communities during wet and dry summers

Questions: What impact do a wet and a dry growing season have on CO₂ dynamics of mire plant communities along a primary succession gradient from the initiation stage to the bog stage? Location: Mires on a land uplift coast, Finland. Methods: We measured CO₂ dynamics and vascular plant green area development on five mires that form a sequence of mire succession. TWINSPAN was used to define successional mire plant communities and regression analyses were used to explore the temporal variation in CO₂ dynamics of the communities. Results: CO₂ dynamics of successional plant communities reacted differently to a wet and a dry growing season. The net CO₂ uptake rate of the earlier successional communities decreased in the dry growing season due to a decrease in photosynthesizing leaf area. Concurrently, CO₂ uptake of the later successional communities moderately increased or did not change. Generally, the difference in net ecosystem exchange (NEE) between a dry and a wet year resulted from the altered rate of gross photosynthesis (P_G) rather than ecosystem respiration (R_E). Conclusions: Critical factors for the more stable carbon (C) gas dynamics in the later stages of mire succession were (1) higher autogenic control of the physical environment and (2) an increase in the number of factors regulating the P_G rate. These factors may buffer mire ecosystems (in terms of the C sink function) from extreme and unfavourable variations in environmental conditions.     

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.     

Multiple Recruitment Limitation Causes Arrested Succession in Mediterranean Cork Oak Systems

Lack of tree regeneration and persistency of species-poor shrublands represent a growing problem across Mediterranean evergreen oak forests. What constrains forest regeneration is poorly understood, and restoration attempts have been largely unsuccessful. We assessed the contribution of four different mechanisms of tree recruitment limitation (that is, source, dispersal, germination, and establishment) in a cork oak (Quercus suber) system in southern Portugal. Using a combination of field studies and experiments, we quantified seed production, seed removal and dispersal, seed survival and germination, seedling establishment and survival, as well as cork oak natural regeneration for the three dominant vegetation types in this system (Cistus ladanifer shrubland, oak forest, and oak savanna). We found that all four forms of cork oak recruitment limitation were significantly more severe in shrublands than in oak forests and savannas, so that oak seedling recruitment in shrubland was impeded in multiple ways. Our results explain why transitions from shrublands to oak savannas and forests are extremely difficult, and that the release from arrested succession in this system requires the simultaneous relief of multiple constraints on recruitment limitation in the early life history of oaks. These results have important implications for the restoration and conservation of Mediterranean oak systems.     

Tropical Montane Forest Restoration in Costa Rica: Overcoming Barriers to Dispersal and Establishment

Tropical forests are being cleared at an alarming rate although our understanding of their ecology is limited. It is therefore essential to design restoration experiments that both further our basic knowledge of tropical ecology and inform management strategies to facilitate recovery of these ecosystems. Here we synthesize the results of research on tropical montane forest recovery in abandoned pasture in Costa Rica to address the following questions: (1) What factors limit tropical forest recovery in abandoned pasture? and (2) How can we use this information to design strategies to facilitate ecosystem recovery? Our results indicate that a number of factors impede tropical forest recovery in abandoned pasture land. The most important barriers are lack of dispersal of forest seeds and seedling competition with pasture grasses. High seed predation, low seed germination, lack of nutrients, high light intensity, and rabbit herbivory also affect recovery. Successful strategies to facilitate recovery in abandoned pastures must simultaneously overcome numerous obstacles. Our research shows that establishment of woody species, either native tree seedlings or early‐successional shrubs, can be successful in facilitating recovery, by enhancing seed dispersal and shading out pasture grasses. On the contrary, bird perching structures alone are not an effective strategy, because they only serve to enhance seed dispersal but do not reduce grass cover. Remnant pasture trees can serve as foci of natural recovery and may enhance growth of planted seedlings. Our results highlight the importance of: (1) understanding the basic biology of an ecosystem to design effective restoration strategies; (2) comparing results across a range of sites to determine which restoration strategies are most generally useful; and (3) considering where best to allocate efforts in large‐scale restoration projects.     

Seed size and dispersal potential of Acer rubrum (Aceraceae) samaras produced by populations in early and late successional environments

In order to determine if red maple dispersal potential or seed size change during secondary succession, samaras were collected from five populations located in early successional environments and five populations located in late successional environments. Wing loading ratios (samara mass—mg/samara area—cm²), which are inversely proportional to dispersal ability, were computed for all samaras, and seeds were excised from each samara and weighed. Samaras from the early successional red maples showed slightly but significantly lower wing loading ratios than those from the late successional environments. This result corresponds with the conclusions reached by several theoretical investigations of seed dispersal evolution that predict that recently founded populations will show greater dispersal abilities than more established populations. The earlier successional populations had slightly heavier seeds than the later successional populations, which suggests that the changes in community composition and dynamics that occur during this successional sequence do not select for heavier seeds in older red maple populations. Coefficients of variation for wing loading and seed size showed no consistent trends with successional stage, which indicates that variation in these characters does not decrease as succession proceeds.     

Seed rain,seedling establishment and clonal growth strategies on a glacier foreland

Abstract. Seed production, composition of the seed rain, germination, and seedling mortality, as well as vegetative growth characteristics of common pioneer plant species were studied on the foreland of the retreating Morteratsch glacier in the Swiss Alps. The frequency of diaspores trapped in different successional stages was related to their dispersal mode and was highly skewed towards a few species. Plenty of diaspores well adapted for dispersal by wind are a precondition for the most important pioneer species. Seeds from all pioneer species investigated had a good germination success, provided that the moisture content of the soil was high enough. However, requirement for seedling establishment differed among sites of increasing terrain age and among species. Only specialized pioneers such as Cerastium pedunulatum, Linaria alpina, Oxyria digyna and Saxifraga aizoides tolerate the cold and moist conditions near the glacier. However, these species are restricted to early successional stages. Seedlings of Epilobium fleischeri are affected not only by the cold and moist conditions near the glacier but also by moderately dry conditions on older sites. Availability of safe sites becomes crucial for most species with increasing age of sites and with drier conditions. Most species playing a dominant role during early succession and persisting during later successional stages have a distinctive ability to spread clonally and have a growth form with more or less widely spaced ramets: Achillea moschata. Cerastium pedunculatum. Epilobium fleischeri and Hieraium staticifolium. The growth strategy and demography of the clonal E. fleischeri is presented as an example. The life cycle of this species is characterized in succession by (1) the colonization of safe sites by small seeds adapted for wind dispersal, (2) horizontal spread by clonal growth, and (3) the persistence through phenotypic morphological plasticity in later successional stages. Seedling establishment and clonal growth are thus complementary mechanisms in plant succession on recently deglaciated terrain.     

Recruitment limitation of forest communities in a degraded Mediterranean landscape

Question: How does habitat degradation affect recruitment limitation and its components (seed limitation versus establishment limitation) of woody plant communities in a Mediterranean landscape? Location: 1600‐1900 m a.s.l. in the Sierra Nevada National Park, southern Spain. The landscape is a mosaic composed of native forest and two degraded landscape units: reforestation stands and shrubland. Methods: We evaluated fruit production, seed rain, seedling emergence and seedling survival in two consecutive years with contrasting rainfall patterns. Seed and seedling data were used to calculate values of seed and establishment limitation. Results: In general, the woody community was both severely seed‐ and establishment‐limited. Species were less seed‐limited in the landscape units with higher adult density (i.e. shrub species in shrubland, Pinus spp. in reforestation stands). In contrast, degradation did not exacerbate establishment limitation, which was severe in all landscape units. This general pattern was modulated by the biogeographical distribution, dispersal type, and life form of the species. Boreo‐alpine species were more limited in establishment than species with a typical Mediterranean distribution. Zoochorous species were less seed‐limited in the landscape units preferred by dispersers (i.e. native forest). Tree species were more establishment‐limited than shrub species, irrespective of the landscape unit. Seed limitation, and especially establishment limitation, varied among years, with establishment being almost nil in the very dry year. Conclusion: In the case of Mediterranean landscapes, when degradation from human impact involves a reduction in the adult abundance of the woody plant community (trees and shrubs), seed limitation increases, although establishment limitation is generally high in all landscape units, especially for boreo‐alpine species. Conservation and restoration strategies should take into account our results showing that tree species were unable to recruit in an extremely dry year, because more aridity is expected under a climatic change scenario in Mediterranean ecosystems.     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.     

The role of land-use history in driving successional pathways and its implications for the restoration of tropical forests

Secondary forests are increasingly important components of human-modified landscapes in the tropics. Successional pathways, however, can vary enormously across and within landscapes, with divergent regrowth rates, vegetation structure and species composition. While climatic and edaphic conditions drive variations across regions, land-use history plays a central role in driving alternative successional pathways within human-modified landscapes. How land use affects succession depends on its intensity, spatial extent, frequency, duration and management practices, and is mediated by a complex combination of mechanisms acting on different ecosystem components and at different spatial and temporal scales. We review the literature aiming to provide a comprehensive understanding of the mechanisms underlying the long-lasting effects of land use on tropical forest succession and to discuss its implications for forest restoration. We organize it following a framework based on the hierarchical model of succession and ecological filtering theory. This review shows that our knowledge is mostly derived from studies in Neotropical forests regenerating after abandonment of shifting cultivation or pasture systems. Vegetation is the ecological component assessed most often. Little is known regarding how the recovery of belowground processes and microbiota communities is affected by previous land-use history. In published studies, land-use history has been mostly characterized by type, without discrimination of intensity, extent, duration or frequency. We compile and discuss the metrics used to describe land-use history, aiming to facilitate future studies. The literature shows that (i) species availability to succession is affected by transformations in the landscape that affect dispersal, and by management practices and seed predation, which affect the composition and diversity of propagules on site. Once a species successfully reaches an abandoned field, its establishment and performance are dependent on resistance to management practices, tolerance to (modified) soil conditions, herbivory, competition with weeds and invasive species, and facilitation by remnant trees. (ii) Structural and compositional divergences at early stages of succession remain for decades, suggesting that early communities play an important role in governing further ecosystem functioning and processes during succession. Management interventions at early stages could help enhance recovery rates and manipulate successional pathways. (iii) The combination of local and landscape conditions defines the limitations to succession and therefore the potential for natural regeneration to restore ecosystem properties effectively. The knowledge summarized here could enable the identification of conditions in which natural regeneration could efficiently promote forest restoration, and where specific management practices are required to foster succession. Finally, characterization of the landscape context and previous land-use history is essential to understand the limitations to succession and therefore to define cost-effective restoration strategies. Advancing knowledge on these two aspects is key for finding generalizable relations that will increase the predictability of succession and the efficiency of forest restoration under different landscape contexts.     

动物传播者对植物更新的促进与限制

在动物 植物的关系网络中,传播者对植物更新具有促进与限制两方面作用.本文从种群尺度总结了传播者取食、空间利用等行为对植物更新的影响;从群落尺度分析了多种传播者传播有效性对植物更新的促进与限制.传播者对食物的处理方式决定了种子的命运,且具有明显的种间差异;植物在传播者食谱中的地位亦决定其更新的成败,成为动物偏好的食物可助其摆脱森林破碎化等不利事件的影响.动物的空间行为可导致种子命运发生改变.传播者移动距离能否逃逸同种成树对种子的距离限制,影响种子的更新命运;动物偏好的适宜生境与适于植物萌发生境的空间一致性程度影响传播者传播的成效.有效传播者的非冗余性促使种子传播网络更稳定,利于植物更新;无效传播直接限制植物更新,但为其他植物定殖提供了可用空间.今后应将传播者行为融入植物种群更新研究,而从生态系统服务角度揭示传播者在植被恢复的作用应是未来恢复生态学研究的重点.     

Soil seed bank and driftline composition along a successional gradient on a temperate salt marsh

Abstract. This study focuses on the relationship between vegetation succession and soil seed bank composition on the Schiermonnikoog (The Netherlands) salt marsh over 100 yr. The importance of driftline material in seed dispersal and the relationship with succession is also investigated. The results indicate that the majority of species have a transient or short‐term seed persistent bank. Seeds of most species are able to float over the salt marsh and become concentrated in the driftline higher up the marsh. After plants have established a seed bank forms, which disappears when vegetation is replaced by later‐successional species. Exceptions are Spergularia mar‐itima, which is still present in the seed bank of late successional stages, and Juncus gerardi and Glaux maritima, which appear in the seed bank of early successional stages, but are absent in the vegetation. Based on the results of this study constraints and possibilities for salt‐marsh restoration by de‐embankment are discussed.