The role of spontaneous vegetation succession in ecosystem restoration: A perspective

Abstract. The paper summarizes ideas which were discussed during the ‘Spontaneous Succession in Ecosystem Restoration’ conference and elaborated through further discussion among the authors. It seeks to promote the integration of scientific knowledge on spontaneous vegetation succession into restoration programs. A scheme illustrating how knowledge of spontaneous succession may be applied to restoration is presented, and perspectives and possible future research on using spontaneous vegetation succession in ecosystem restoration are proposed. It is concluded that when implementing spontaneous succession for ecological restoration the following points must be considered: setting clear aims; evaluation of environmental site conditions; deciding whether spontaneous succession is an appropriate way to achieve the aims; prediction of successional development; monitoring of the results. The need for interdisciplinary approaches and communication between scientists, engineers and decision‐makers is emphasized.     

Spontaneous succession in limestone quarries as an effective restoration tool for endangered arthropods and plants

1. The view of post-mining sites is rapidly changing among ecologists and conservationists, as sensitive restoration using spontaneous succession may turn such sites into biodiversity refuges in human-exploited regions. However, technical reclamation, consisting of covering the sites by topsoil, sowing fast-growing herb mixtures and planting trees, is still commonly adopted. Until now, no multi-taxa study has compared technically reclaimed sites and sites left with spontaneous succession.
2. We sampled communities of vascular plants and 10 arthropod groups in technically reclaimed and spontaneously restored plots in limestone quarries in the Bohemian Karst, Czech Republic. For comparison, we used paired t -tests and multivariate methods, emphasizing red-list status and habitat specialization of individual species.
3. We recorded 692 species of target taxa, with a high proportion of red-listed (10%) and xeric specialist (14%) species, corroborating the great conservation potential of the quarries.
4. Spontaneously restored post-mining sites did not differ in species richness from the technical reclaimed sites but they supported more rare species. The microhabitat cover of leaf litter, herbs and moss, were all directly influenced by the addition of topsoil during reclamation.
5. Synthesis and applications. Our results show that the high conservation potential of limestone quarries could be realized by allowing succession to progress spontaneously with minimal intervention. Given the threat to semi-natural sparsely vegetated habitats in many regions, active restoration measures at post-mining sites should be limited to maintenance of early successional stages, instead of acceleration of succession.     

Effects of Montane Heathland Restoration on Leafhopper Assemblages (Insecta: Auchenorrhyncha)

At the beginning of the 20th century, many montane heathlands were abandoned and became subject to natural succession or afforestation by humans. Thus, the formerly large montane heathlands slowly degraded into small and isolated patches. In this study, we evaluate the influence of restoration measures on leafhopper (Auchenorrhyncha) assemblages of montane heathland ecosystems in Central Europe. Our analyses comprised three different site types that were adjacent to each other: (1) montane heathlands, (2) restoration sites, and (3) control sites. Leafhoppers showed a clear response to montane heathland restoration. Thus, after 4–5 years since implementation of restoration measurements restoration sites were characterized by the highest species richness. However, detailed analyses of leafhopper diversity, species composition, and environmental parameters on the three site types revealed that restoration sites were rather similar to control sites and significantly differing from montane heathlands. We conclude that leafhoppers are excellent bioindicators for restoration measurements because they reflected environmental differences between the three site types. Restoration measurements might only be a useful instrument to promote typical montane heathland leafhopper communities in the long run. Colonization by leafhoppers is, however, dependent on many different factors such as leafhopper mobility, vegetation structure, microclimate, and the establishment of ericaceous dwarf shrubs. Practitioners should establish a management regime (grazing and sod‐cutting) that creates a mosaic of different habitat structures and increases typical heathland vegetation, thus, favoring the colonization of typical heathland leafhoppers.     

Secondary succession on sandy old‐fields in Hungary

Abstract. Development of semi‐natural vegetation has recently been a primary concern of restoration efforts. A primary management question is whether active intervention is required or spontaneous secondary succession could suffice. We studied 54 old‐fields in central Hungary, which differed in time since abandonment but which had similar environmental conditions and management histories. The sites were grouped into four age groups according to the time elapsed since cultivation abandonment: 1–5, 6–10, 11–23 and 24–33 yr. In each old‐field we recorded the species and estimated their abundances. We grouped species in two ways: according to life form (annuals, biennials, perennials, woody plants) and according to coenological behaviour (weeds, sand and steppe generalists, specialists). We analysed the changes in species number and abundance in these categories as a function of site age. Contrary to other successional studies, the total number of species did not change significantly among the four age groups. A significant change was detected between the first two age groups as to life‐form composition. Species number and abundance of annuals decreased, while the perennials and woody plants increased. As to coenological behaviour, species number changed only in the first two age groups, while abundance changed in the first three. Weeds quickly disappeared and specialists established and spread, while the species number and abundance of generalists did not change significantly. We concluded that the basic shifts in species composition are almost completed within 10 yr. Most of the late successional species colonized and weeds disappeared. We conclude that there was no need for active intervention in this system: the spontaneous secondary succession leads to semi‐natural vegetation.     

采石场废弃地的生态重建研究进展

采石场的开采严重破坏了植被和土壤,形成了大量的裸露岩石斜坡,造成宏观景观支离破碎和极端的环境条件,限制了植物的生长。由于自然恢复所需时间长久,人工恢复被广泛应用于采石场废弃地的生态重建。自然演替过程是采石场生态重建的理论基础,自然演替理论可以为人工恢复措施提供指导。植物群落演替的早期阶段,非生物因素起主要作用,随着演替的推移,生物因素的重要性增强。邻近自然植被的土壤和繁殖体通过外力的扩散,对恢复起重要作用。除了非生物和其他的限制,先到达恢复地的物种竞争能力的变化能决定了演替过程。演替过程中的干扰因素往往成为演替重要的驱动力。裸露岩石斜坡的物理稳定性对植被恢复有重要影响,有机废物的使用和施肥可以影响恢复演替的方向和生物多样性。播种一定的植物能够改变恢复演替方向,加速演替过程。乡土物种适应了当地气候,能够促进演替。随着修复时间的延长,土壤有机质含量,植被覆盖度和物种丰富度不断增加,土壤微生物生物量随之增加。开展不同地区采石场植物种类的选育、研究乡土物种的功能特性、土壤微生物群落和酶的变化、植被演替过程的定位研究、植物种间的竞争关系、自然演替和人工恢复的比较研究、探索经济高效的采石场生态重建方法是未来的研究方向。     

Increasing intraspecific facilitation in exposed environments: consistent results from mountain birch populations in two subarctic stress gradients

The stress‐gradient hypothesis predicts that the importance of facilitation relative to competition should increase with increasing stress. The hypothesis has received support from several environments, but multi‐gradient studies on the generality of the hypothesis are exceptionally rare. A within‐species experiment with mountain birch Betula pubescens subsp. czerepanovii was conducted to test the hypothesis in the extreme ends of two subarctic stress gradients (elevation and seashore) in the Kola Peninsula, northwestern Russia. The high stress sites were characterized by strong winds, temperature extremes and potentially drought. The negative effects of abiotic stress on the study seedlings were verified from performance characteristics. Effects of adult hosts as well as seedling–seedlings interactions were studied. Positive host–seedling interactions dominated in each study site, and three out of four performance variables indicated stronger positive net effects in the high stress sites. In the seashore gradient also seedling survival gave similar interpretations. Also a temporal shift towards host–seedling competition was detected in a low stress site after two study years. In seedling–seedling interactions competition dominated, but the effects were weak, likely due to the ‘noise’ caused by genetic and environmental factors. Our results support the stress‐gradient hypothesis and its generality in subarctic environments, as the interpretations were similar for both stress gradients and several fitness‐related variables. The temporal variation in host–seedling interactions and the difference between host–seedling and seedling–seedling effects hint on size‐dependency of plant interactions: facilitation might dominate when the benefactor is substantially larger than the beneficiary, while competition may be stronger when the plants are of similar size and developmental status.     

科尔沁沙地植被恢复系列上群落演替与物种多样性的恢复动态

通过对科尔沁沙地植被恢复系列上不同阶段的群落取样,研究了植被恢复过程中群落物种组成、物种多样性的变化过程.在演替时间分别为1年、3年、5年、12年、20年和30年的群落中,物种丰富度分别为7、11、17、14、28和30种,而物种多样性指数分别为1.458 8、2.610 0、3.108 4、1.696 8、3.738 6、3.639 6.在生活型结构上,一年生植物的种类数量占绝对优势,但随着演替进展,多年生植物的种类数量增加且单种植物的优势度超过了一年生植物,表明多年生植物在群落功能维持中占据重要地位.随演替进展,来自不同科属的植物增加.藜科植物在演替过程中种类数量较为稳定,特别是在演替早期藜科植物占有很高的优势度.禾本科种类随演替进展不断增加,且在群落中的优势度逐渐上升.总体上,随演替进展群落种类组成与物种多样性增加,群落生态优势度下降,而均匀度增加,群落趋向稳定.     

Effects of sample distribution along gradients on eigenvector ordination

In general, disproportionately heavy sampling of the ends of a gradient increases the interpretability of eigenvector ordinations. More specifically, correspondence analysis (CA) and detrended correspondence analysis (DCA) best reproduce the original positions of samples in simulated coenoclines when samples are clustered toward the ends of the axis. Principal components analysis (PCA) reproduces the original sample positions less well than either CA or DCA and shows no improvement as samples are increasingly clustered toward the ends of the axis. PCA and CA show less curvature of one dimensional data into the second axis when sampling favors the ends of the axis.     

Do not neglect surroundings in restoration of disturbed sites

We examined the influence of surroundings, i.e. proximity effects, on the course of spontaneous vegetation succession using two data sets. In the first data set, we tested the effects of surrounding vegetation (woodlands, wetlands, grasslands, and synanthropic) on succession in various disturbed habitats in the Czech Republic by comparing successional sites with more natural vegetation within 100 m and at 1 km from each site. The habitats included old fields, gravel‐sand pits, spoil heaps from black coal mining, industrially extracted peatlands, and acidic stone quarries. We found that, with the exception of wetlands, the influence of the vegetation types on seral vegetation was nearly always significant using marginal and partial Canonical Correspondence Analyses. In the second data set, which included 27 limestone quarries, we compared species lists outside (up to 100 m apart) and inside the quarries using Detrended Correspondence Analysis and the Sørensen similarity index. We found much higher species similarity between outside and inside particular limestone quarries than among the quarries themselves and among their surroundings, which also indicates that the seral vegetation is decisively influenced by the surroundings. We argue that restoration ecologists should carefully consider the nature of the surroundings of disturbed sites because of its profound impact on restoration processes. They should conduct inventories and prescribe some restoration measures not only inside a restored site, but also in its surroundings.     

Vegetation succession on reclaimed coal wastes in Spain: the influence of soil and environmental factors

Question: How is vegetation succession on coal mine wastes under a Mediterranean climate affected by the restoration method used (topsoil addition or not)? How are plant successional processes influenced by local landscape and soil factors? Location: Reclaimed coal mines in the north of Palencia province, northern Spain (42°47′‐42°50′ N, 4°32′‐4°53′ W). Methods: In Jun–Jul 2008, vascular plant species cover was monitored in 31 coal mines. The mines, which had been restored using two restoration methods (topsoil addition or not), comprised a chronosequence of different ages from 1 to 40 yr since restoration started. Soil and environmental factors at each mine were monitored and related to species cover using a combination of ordination methods and Huisman–Olff–Fresco modeling. Results: Plant succession was affected by restoration method . Where topsoil was added, succession was influenced by age since restoration and soil pH. Where no topsoil was added, soil factors seem to arrest succession. Vegetation composition on topsoiled sites showed a gradient with age, from the youngest, with early colonizing species, to oldest, with an increase in woody species. Vegetation on non‐topsoiled sites comprised mainly early‐successional species. Response to age and pH of 37 species found on topsoiled mines is described. Conclusions: Restoration of coal mines under this Mediterranean climate can be relatively fast if topsoil is added, with a native shrub community developing after 15 yr. However, if topsoil is not used, it takes more than 40 yr. For topsoiled mines, the species found in the different successional stages were identified, and their tolerance to soil pH was derived. This information will assist future restoration projects in the area.     

演替理论与植被恢复

《昆明-蒙特利尔全球生物多样性框架》提出要高质量保护和恢复各30%的土地,最大化地实现保护生物多样性和缓解气候变化的目标,而演替理论和植被恢复可以为实现30%的保护和恢复目标服务。演替理论是植被生态学中的核心理论,演替是指在一个地点上由一群不同物种组成的生命体的结构或组成随时间而变化的过程; 植被恢复是以植物种植、配置为主,恢复或重建植物群落或天然更新恢复植物群落的过程,植被恢复是生态系统结构和功能从简单到复杂、从低级向高级变化的过程,最终目的是建立健康稳定的植物群落。演替是植被恢复的基础,植被恢复被视为对演替过程的操纵,以达到恢复受损植被生态系统的目标。演替理论可以指导植被恢复,而植被恢复对演替理论的发展有益。演替按裸地性质可以分为原生演替和次生演替,有研究建议将恢复过程视为第三演替,这将有助于理解通过人为干预促进植被恢复成功的管理选择,特别是通过强调退化生态系统中的环境和生物遗存的管理选择。此外,该文还提出了植被恢复理论和演替理论未来可能重点关注的科学和技术问题。     

Spontaneous succession in Central‐European man‐made habitats: What information can be used in restoration practice?

Abstract. The results of previous studies concerning spontaneous vegetation succession in various man‐made or disturbed habitats in Central Europe are discussed in relation to restoration ecology. An attempt is made to answer the main questions which are important to restoration programs. These concern the rate and direction of succession; participation of target species and communities, woody species, ruderals and aliens; arresting or diverting succession; changes in species diversity; variability of successional pathways and the role of abiotic environmental conditions. The great potential for using successional theory in restoration programs is emphasized.     

A call for applying trophic structure in ecological restoration

Ecological restoration projects have traditionally focused on vegetation as both a means (seeding, planting, and substrate amendments) and ends (success based upon primary productivity and vegetation diversity). This vegetation‐centric approach to ecological restoration stems from an historic emphasis on esthetics and cost but provides a limited measure of total ecosystem functioning and overlooks alternative ways to achieve current and future restoration targets. We advocate a shift to planning beyond the plant community and toward the physical and biological components necessary to initiate autogenic recovery, then guiding this process through the timely introduction of top predators and environmental modifications such as soil amendments and physical structures for animal nesting and refugia.     

Biological Soil Crust Rehabilitation in Theory and Practice: An Underexploited Opportunity

Biological soil crusts (BSCs) are ubiquitous lichen–bryophyte microbial communities, which are critical structural and functional components of many ecosystems. However, BSCs are rarely addressed in the restoration literature. The purposes of this review were to examine the ecological roles BSCs play in succession models, the backbone of restoration theory, and to discuss the practical aspects of rehabilitating BSCs to disturbed ecosystems. Most evidence indicates that BSCs facilitate succession to later seres, suggesting that assisted recovery of BSCs could speed up succession. Because BSCs are ecosystem engineers in high abiotic stress systems, loss of BSCs may be synonymous with crossing degradation thresholds. However, assisted recovery of BSCs may allow a transition from a degraded steady state to a more desired alternative steady state. In practice, BSC rehabilitation has three major components: (1) establishment of goals; (2) selection and implementation of rehabilitation techniques; and (3) monitoring. Statistical predictive modeling is a useful method for estimating the potential BSC condition of a rehabilitation site. Various rehabilitation techniques attempt to correct, in decreasing order of difficulty, active soil erosion (e.g., stabilization techniques), resource deficiencies (e.g., moisture and nutrient augmentation), or BSC propagule scarcity (e.g., inoculation). Success will probably be contingent on prior evaluation of site conditions and accurate identification of constraints to BSC reestablishment. Rehabilitation of BSCs is attainable and may be required in the recovery of some ecosystems. The strong influence that BSCs exert on ecosystems is an underexploited opportunity for restorationists to return disturbed ecosystems to a desirable trajectory.     

Community maturity, species saturation and the variant diversity-productivity relationships in grasslands

Detailed knowledge of the relationship between plant diversity and productivity is critical for advancing our understanding of ecosystem functioning and for achieving success in habitat restoration efforts. However, effects and interactions of diversity, succession and biotic invasions on productivity remain elusive. We studied newly established communities in relation to preexisting homogeneous vegetation invaded by exotic plants in the northern Great Plains, USA, at four study sites for 3 years. We observed variant diversity–productivity relationships for the seeded communities (generally positive monotonic at three sites and non-monotonic at the other site) but no relationships for the resident community or the seeded and resident communities combined at all sites and all years. Community richness was enhanced by seeding additional species but productivity was not. The optimal diversity (as indicated by maximum productivity) changed among sites and as the community developed. The findings shed new light on ecosystem functioning of biodiversity under different conditions and have important implications for restoration.     

More Ecology is Needed to Restore Mediterranean Ecosystems: A Reply to Valladares and Gianoli

Valladares and Gianoli (2007) tried to answer a key question, “how much ecology do we need to know to restore Mediterranean ecosystems?” by focusing on (1) plant–plant interactions; (2) environmental heterogeneity and the potential adaptation of transplanted plants; and (3) phenotypic plasticity of the planted species. We consider their choice of topics incomplete and potentially misleading because (1) it is clearly biased toward a narrow set of research topics (phenotypic plasticity, facilitation, and climate change); (2) it assumes that active restoration, and specifically revegetation, is needed; and (3) it conveys a false perception that other basic ecological aspects of Mediterranean ecosystems are sufficiently known. Instead, we review the current knowledge on seed dispersal, succession, and ecosystem functioning for Mediterranean ecosystems. We argue that decades of research on these topics have yielded few practical guidelines for restoration, something that needs to be urgently corrected. First, the current “establishment limitation paradigm” for plant recruitment does not acknowledge the role of dispersal limitation at large spatial scales. More attention should be paid to nucleation processes and directed seed dispersal mediated by animals. Second, studies of vegetation dynamics and succession in the Mediterranean have led to an overly simplistic view of successional dynamics. How fast and deterministic succession is remains mostly unexplored; long‐term monitoring of successional dynamics at different spatial scales is urgently needed. Third, information on the functional status of Mediterranean ecosystems is required to identify processes hindering natural recovery after disturbances and to set priorities on the areas and ecosystem components to be restored.     

Grassland restoration on former croplands in Europe: an assessment of applicability of techniques and costs

Grasslands used to be vital landscape elements throughout Europe. Nowadays, the area of grasslands is dramatically reduced, especially in industrial countries. Grassland restoration is widely applied to increase the naturalness of the landscape and preserve biodiversity. We reviewed the most frequently used restoration techniques (spontaneous succession, sowing seed mixtures, transfer of plant material, topsoil removal and transfer) and techniques used to improve species richness (planting, grazing and mowing) to recover natural-like grasslands from ex-arable lands. We focus on the usefulness of methods in restoring biodiversity, their practical feasibility and costs. We conclude that the success of each technique depends on the site conditions, history, availability of propagules and/or donor sites, and on the budget and time available for restoration. Spontaneous succession can be an option for restoration when no rapid result is expected, and is likely to lead to the target in areas with high availability of propagules. Sowing low-diversity seed mixtures is recommended when we aim at to create basic grassland vegetation in large areas and/or in a short time. The compilation of high-diversity seed mixtures for large sites is rather difficult and expensive; thus, it may be applied rather on smaller areas. We recommend combining the two kinds of seed sowing methods by sowing low-diversity mixtures in a large area and high-diversity mixtures in small blocks to create species-rich source patches for the spontaneous colonization of nearby areas. When proper local hay sources are available, the restoration with plant material transfer can be a fast and effective method for restoration.     

内蒙古草原退化群落恢复演替的研究II. 恢复演替时间进程的分析

 根据内蒙古典型草原地带的羊草+大针茅草原退化变型一冷蒿群落封育12年(1983—1994)的动态监测数据进行分析,对群落恢复演替轨迹取得以下认识： 1．依据群落优势种的更替及主分量分析结果可将恢复演替过程划分为冷蒿优势阶段、冷蒿+冰草阶段、冰草优势阶段、羊草优势阶段。 2;退化草原群落在恢复演替过程中,群落生产力的变化表现出阶梯式跃变和亚稳态阶面相间的特点。第一次跃变发生在1984年,上升到第二个阶面,第二次跃变发生在1990年,进入了第三个阶面,已接近于原生群落的生产力。 3．群落生产力与水资源量的关系因恢复演替阶段不同而异。第一亚稳态时期,群落地上现存生物量大体处于166g·m-2的水平上,生长季降水量达176mm以上时,增加降水对群落生产力的提高不发生显著影响。第二亚稳态时期,群落生物量与降水量之间的相关性显著。可推算出群落于物质生产用水量介于1.1～1.6mm·g-1之间。此值在1.1mm·g-1时,群落对水资源的利用效率最高,而在1.6mm·g-1时群落生物量达到最大值。 4．在恢复演替进程中,群落密度的位点常数约为271.5株·m-2,循此常数上下波动,表现出拥挤与稀疏交替发生的过程,构成了恢复演替的节奏性变化。群落生物量的跃变与亚稳态的形成,以及群落密度的拥挤与稀疏交替作用是群落恢复演替的内在机制。恢复演替的速度,到第10年发生了1.78个半变的生态距离。5．草原退化群落恢复演替过程中,按照其节奏性及生产力跃变与亚稳态的规律,调控放牧利用强度或采取技术措施,调节群落拥挤和稀疏的交替过程可加速恢复演替进程。     

长期封育演替下典型草原植物群落生产力与多样性关系

封育是退化草地的重要恢复措施, 理解长期封育过程中草地群落生产力和植物多样性变化特征及两者间关系, 有助于草地植被的恢复管理与利用。该研究依托宁夏云雾山国家级自然保护区典型草原长期封育演替梯度, 选择持续放牧、封育9年、26年和34年的草地群落作为研究对象, 分析其地上生产力、物种多样性和功能多样性的变化特征及内在联系。结果表明, 封育显著提高典型草原植物群落的地上生产力、凋落物生物量、功能丰富度和功能离散度, 未改变草地群落的物种丰富度、Shannon-Wiener指数和功能均匀度, 但Simpson优势度指数和Pielou均匀度指数在长期封育(34年)后显著下降。此外, 封育对不同植物群落加权平均功能性状的影响存在差异。随机森林模型和方差分解结果显示, 群落加权平均功能性状对封育草地群落地上生产力变异的解释度高达70.70%, 其中植株高度是最主要的解释因子; 功能多样性的解释度为36.86%, 主要由功能丰富度贡献; 而物种多样性的解释度仅为14.72%。由此可见, 植物功能性状和功能多样性对草地群落地上生产力的贡献远高于物种多样性, 建议将其纳入植物群落恢复演替动态研究, 以便全面了解植物多样性与生态系统功能的关系, 为更好地实现生态恢复目标奠定基础。     

The Effects of Peatland Restoration on Water‐Table Depth,Elemental Concentrations,and Vegetation: 10 Years of Changes

We studied the effects of restoration on water‐table depth (WTD), element concentrations of peat and vegetation composition of peatlands drained for forestry in southern Finland. The restoration aimed to return the trajectory of vegetation succession toward that of undisturbed systems through the blockage of ditches and the removal of trees. Permanent plots established on a bog and a fen were sampled 1 year before, and 1, 2, 3, and 10 years after the restoration. The restoration resulted in a long‐term rise of the water‐table in both peatlands. Ten years after restoration, the mineral element concentrations (Ca, K, Mg, Mn, and P) of peat corresponded to those reported from comparable pristine peatlands. In particular, the increase of K and Mn concentrations at both sites suggests the recovery of ecosystem functionality in terms of nutrient cycling between peat and plants. The restoration resulted in the succession of plant communities toward the targeted peatland vegetation of wetter condition at both sites. This was evident from the decreased abundance of species benefiting from drainage and the corresponding increase of peatland species. However, many species typical of pristine peatlands were missing 10 years after restoration. We conclude that the restoration led to a reversal of the effects of drainage in vegetation and studied habitat conditions. However, due to the slow recovery of peatland ecosystems and the possibility that certain failures in the restoration measures may become apparent only after extended time periods, long‐term monitoring is needed to determine whether the goals of restoration will be met.