Rate of succession in restored wetlands and the role of site context

Question: Are changes in plant species composition, functional group composition and rates of species turnover consistent among early successional wetlands, and what is the role of landscape context in determining the rate of succession? Location: Twenty‐four restored wetlands in Illinois, USA. Methods: We use 4 years of vegetation sampling data from each site to describe successional trends and rates of species turnover in wetlands. We quantify: (1) the rate at which composition changes from early‐successional to late‐successional species and functional groups, as indicated by site movement in ordination space over time, and (2) the rate of change in the colonization and local extinction of individual species. We correlate the pace of succession to site area, isolation and surrounding land cover. Results: Some commonalities in successional trends were evident among sites. Annual species were replaced by clonal perennials, and colonization rates declined over time. However, differences among sites outweighed site age in determining species composition, and the pace of succession was influenced by a site's landscape setting. Rates of species turnover were higher in smaller wetlands. In addition, wetlands in agricultural landscapes underwent succession more rapidly, as indicated by a rapid increase in dominance by late‐successional plants. Conclusions: Although the outcome of plant community succession in restored wetlands was somewhat predictable, species composition and the pace of succession varied among sites. The ability of restoration practitioners to accelerate succession through active manipulation may be contingent upon landscape context.     

Successional leafhopper assemblages: Pattern and process

Auchenorrhyncha (leafhopper) faunas of a series of experimental plots of different successional age were recorded by vacuum sampling. There were clear successional trends in the assemblages. The life-history strategy of the dominant species changed during succession; the mobile and multivoltine or bivoltine species, which overwinter as adults or nymphs, are replaced by less active, univoltine or bivoltine species which overwinter as eggs. Leafhopper assemblages clearly responded to the changes in plant species composition and vegetation structure during succession; however, disentangling the relative importance of the various factors requires further experimental manipulation.     

不同演替阶段鼢鼠土丘群落植物多样性变化研究

用空间序列代替时间序列的方法对高寒草甸不同演替阶段高原鼢鼠土丘植物群落的物种组成和多样性变化进行了研究．结果表明,不同演替阶段鼠丘植物群落的物种组成及外貌特征与原生植被(对照)之间存在较大差异．在演替的早期阶段,r对策者如萼果香薷、灰绿藜、鹅绒委陵菜、细叶亚菊等演替先锋种在群落中占相对重要地位．随着演替的进展,k对策者如长毛风毛菊、垂穗披碱草、甘肃嵩草、线叶嵩草等在群落中的比例增加．α多样性分析表明,随着演替的进展,群落物种丰富度指数(O)显著增加,其排列顺序为：阶段1<阶段2<阶段3<阶段4<原生植被;均匀度指数(Pielou均匀度指数)的变化趋势与丰富度指数相同;多样性指数(Simpson指数D和Shannon-．Wiener指数H’)按群落的演替梯度呈增加趋势．卢多样性分析表明,阶段1与原生植被及阶段1与阶段4植物群落物种组成的相似系数最小,为0．18;阶段1与阶段2植物群落物种组成的相似系数最大,为0．62．同时,群落生活型功能群组成也随鼠丘的演替进展而发生变化,反映出不同演替阶段的鼠丘植物群落和微生境都发生了改变．     

Secondary Succession and Natural Habitat Restoration in Abandoned Rice Fields of Central Korea

Floristic composition and soil characteristics (moisture, pH, nutrient contents) in abandoned upland rice paddies of different ages were analyzed to clarify the regenerative aspects of succession as a tool for habitat restoration. The study sites represented five seral stages: newly abandoned paddy fields; successional paddy fields abandoned for 3, 7, and 10 years; and a 50‐year‐old Alnus japonica forest. A vegetation sere was apparent in changes of dominant plant species in the order Alopecurus aequalis var. amurensis (annual grass), Aneilema keisak (annual forb), Juncus effusus var. decipiens (rush), Salix koriyanagi (willow), and Alnus japonica (alder) communities. These temporal stages resemble the spatial zonation of vegetation in local riparian floodplain ecosystems, indicating a hydrosere, with soil moisture decreasing over time. Age distributions and life forms of the dominant plant species support a “tolerance” model of secondary succession, in which the established species persist into later successional stages. Persistence of earlier colonizers led to a net cumulative increase in species richness and a more even distribution of species cover with increasing field age. Between 10 and 50 years, vegetation stabilizes as an alder community. Soil moisture content decreased steadily with paddy field age after an initial rise immediately after their abandonment, whereas pools of organic matter, N, P, K, Ca, and Mg, increased with field age. The pace and direction of recovery of native vegetation and natural soil properties in these abandoned rice paddies resembled classic old field succession, a form of secondary succession that often serves as a template for guiding restoration efforts. Active intervention, in particular dismantling artificial levees, could accelerate the recovery process, but natural habitat recovery generally appears sufficiently robust to achieve “passive” restoration of this rare community without intervention.     

Vegetation succession among and within structural layers following wildfire in managed forests

Question: How does vegetation develop during the initial period following severe wildfire in managed forests? Location: Southwestern Oregon, USA. Methods: In severely burned plantations, dynamics of (1) shrub, herbaceous, and cryptogam richness; (2) cover; (3) topographic, overstory, and site influences were characterized on two contrasting aspects 2 to 4 years following fire. Analysis of variance was used to examine change in structural layer richness and cover over time. Non‐metric multidimensional scaling, multi‐response permutation procedure, and indicator species analysis were used to evaluate changes in community composition over time. Results: Vegetation established rapidly following wildfire in burned plantations, following an initial floristics model of succession among structural layers. Succession within structural layers followed a combination of initial and relay floristic models. Succession occurred simultaneously within and among structural layers following wildfire, but at different rates and with different drivers. Stochastic (fire severity and site history) and deterministic (species life history traits, topography, and pre‐disturbance plant community) factors determined starting points of succession. Multiple successional trajectories were evident in early succession. Conclusions: Mixed conifer forests are resilient to interacting effects of natural and human‐caused disturbances. Predicting the development of vegetation communities following disturbances requires an understanding of the various successional components, such as succession among and within structural layers, and the fire regime. Succession among and within structural layers can follow different successional models and trajectories, occurs at different rates, and is affected by multiple interacting factors.     

黔西北地区不同演替阶段植物群落结构与物种多样性特征

该文采用"空间代替时间"的方法,研究了贵州省威宁县喀斯特地区植被演替过程中的群落结构、物种组成、生活型谱和物种多样性的变化规律。结果表明:(1)该调查共记录到种子植物174种,隶属于52科117属,物种分布较多的有菊科、蔷薇科、禾本科、杜鹃花科、小檗科、唇形科、蓼科。(2)随着植被的正向演替,物种丰富度逐渐增加,群落结构趋于复杂,高位芽植物所占比例逐渐增大。(3)随着植被的恢复,群落层次分化逐渐明显,大径级植株所占比例呈现增加趋势。(4)随着植被的恢复,群落各层次的ShannonWiener多样性指数(H)、Simpson多样性指数(DS)、均匀度指数(J)和Margalef丰富度指数(DM)逐渐增加;不同演替阶段植物群落之间的Srensen相似系数呈现先上升后下降的趋势,Cody指数则表现为逐渐增加的趋势。黔西北地区不同演替阶段植物群落结构和物种多样性不同,建群种和关键种发生了明显变化,不同演替阶段植物群落结构和物种多样性的研究对喀斯特地区植被演替规律的认识和生态恢复具有重要意义。     

Plant succession and rhizosphere microbial communities in a recently deglaciated alpine terrain

This study describes how early and late successional plant species affect soil microorganisms in alpine ecosystems. We quantify the relative importance of plant species and soil properties as determinants of belowground microbial communities. Sixteen plant species were selected from six successional stages (4–14–20–43–75–135 years) within the foreland of the Rotmoosferner glacier, Austria, and at one (reference) site outside the foreland. The size, composition and function of the communities of microorganism in the bulk soil and the rhizosphere were characterized by ninhydrin-reactive nitrogen, phospholipid fatty acids and enzyme activities (β-glucosidase, β-xylosidase, N-acetyl-β-glucosaminidase, leucine aminopeptidase, acid phosphatase, sulphatase). The results show that the microbial data could be grouped according to early (up to 43 years) and late-colonizing plant species (75 or more years). In early succession, no plant species or soil age effect was detected on the microbial biomass, phospholipid fatty acids, or enzyme activity. The rhizosphere microbial community was similar to that in the bulk soil, which in turn was determined by the abiotic environmental conditions. In late succession, improved soil conditions probably mediated plant species effects on the belowground microbial community. The most pronounced rhizosphere effects were attributed to plant species of the 75- and 135-year-old sites. The microbial colonization (size, composition, activity) of the bulk soil predominantly followed changes in vegetation cover, plant life forms and soil organic matter. In summary, the observed successional pattern of the above- and belowground communities provides an example of the facilitation models of primary succession.     

Vegetation development in dune slacks: the role of persistent seed banks

Abstract. The soil seed bank composition was determined at four sites in the dune slack ‘Koegelwieck’ on the Dutch Wadden Sea island of Terschelling. At three different sites in the slack, where sod-cutting experiments down to the mineral sand had been carried out, the established vegetation and seed bank were assessed after 5, 9 and 39 yr of undisturbed development, respectively. In addition, a fourth site in the slack was investigated, where vegetation development had proceeded for 80 yr since plant colonization of bare soil and where nowadays a vegetation dominated by Calamagrostis epigejos and Salix repens occurs. Together these four sites can be regarded as a chronosequence of dune slack formation. Clear time sequences were detected in the seed bank data. Many late successional species showed a significant increase in the number of seeds during the succession. Some of the early successional basiphilous pioneer species such as Anagallis minima, Centaurium littorale, Littorella uniflora, Radiola linoides and Samolus valerandi, showed either a decrease during the time of succession or an optimum in the vegetation while remaining present in the seed bank in low but detectable numbers. They could, therefore, play a role in re-establishment of the vegetation after sod-cutting. One of the target species, Schoenus nigricans, established within a few years after removal of the sod. However, no seeds of this species have been detected in the soil below either of the successional stages. Based on the species disappearance from the established vegetation and based on the independent data of Thompson et al. (1997) an estimation of seed longevity could be made for several Red List species of wet dune slacks.     

Changes in the vegetation of Quercus pubescens woodland after cessation of coppicing and grazing

Abstract. The cessation of coppicing and grazing in Quercus woodlands, with its subsequent changes in the structure, composition and functioning of vegetation communities, is becoming more frequent throughout the Mediterranean Basin. In southern France, we have studied successional changes in Quercus pubescens woodlands by visiting previously studied sites 18 yr later. Changes in vertical structure, species richness, floristic composition, life form and dispersal type were analysed and compared between woodlands that had previously been grazed or ungrazed. Both successions showed a decline in vegetation cover in the 0–25 cm height class and in the height class immediately under the canopy layer, due to oak litter accumulation and tree ageing. In post‐grazing succession, the abandonment of grazing and associated burning has allowed the vegetation cover to increase in the 0.25‐2 m height class. In both successions, grassland species decreased in frequency and forest species increased, a trend which was stronger in undisturbed succession. Species richness decreased with time in the undisturbed succession, but remained stable in the post‐grazing succession mainly because of the slow decline of plants linked to grazing. In undisturbed succession, therophytes and hemicryptophytes decreased. In contrast, therophytes and hemicryptophytes remained stable in post‐grazing succession. In both successions, endozoochorous species (notably Ruscus aculeatus) increased. Plants dispersed by non‐animal vectors decreased in undisturbed succession, but stability was observed in most of the dispersal types in post‐grazing succession. These results showed that a time‐lag existed between undisturbed succession and post‐grazing succession, the latter remaining at a younger stage of successional development due to more recent impact of grazing. However, both successions have converged suggesting that most of the traces of grazing on vegetation will disappear within a few years. The vegetation of these coppices, regardless of the previous grazing regime, will become increasingly similar to the vegetation of undisturbed woodlands. However, their floristic composition will probably never be identical to that of undisturbed woodlands, mainly because of the rarity of these undisturbed woodlands and of the short‐distance dispersal of many forest plant species.     

Primary succession on lava domes on Terceira (Azores)

Questions: Are the vegetation attributes significantly different among lava domes and among geomorphologic units as a result of age and soil features? Are the successional rates equal in all the geomorphologic units of the domes? Are the colonizer species of lava domes totally replaced by other species in the late successional stages? Location: Terceira Island, Azores (Portugal). Methods: Three comparable domes of 240, 370 and 2080 yr old were selected. Data on floristic composition, vegetation bioarea (area occupied by plant species in transects), structure, demography and soil nutrients were collected. Quantitative and qualitative changes along the succession gradient were also analyzed. Results: Vegetation attributes were consistent with the successional stage of each dome in the primary sere; however, the geomorphologic units did not follow the same pattern. The influence of the rates of plant colonization and soil formation are responsible for the decrease of the successional rates from footslopes, to summits, to slopes. The vegetation successionally changes from Juniperus scrub, to Juniperus wood and forest, and there is little species replacement since the similarity in species composition is high between the 3 domes. Conclusions: This is a special type of direct succession that takes place mainly through an increase in biomass and structural complexity. We observed small wooded areas in the fissures of very young domes that are samples of later successional forests — a phenomenon that we call ‘zoom effect’.     

Pathways, mechanisms and predictability of vegetation change during tropical dry forest succession

The development of forest succession theory has been based on studies in temperate and tropical wet forests. As rates and pathways of succession vary with the environment, advances in successional theory and study approaches are challenged by controversies derived from such variation and by the scarcity of studies in other ecosystems. During five years, we studied development pathways and dynamics in a chronosequence spanning from very early to late successional stages (ca. 1–60 years) in a tropical dry forest of Mexico. We (1) contrasted dynamic pathways of change in structure, diversity, and species composition with static, chronosequence-based trends, (2) examined how structure and successional dynamics of guilds of trees shape community change, and (3) assessed the predictability of succession in this system. Forest diversity and structure increased with time but tree density stabilized early in succession. Dynamic pathways matched chronosequence trends. Succession consisted of two tree-dominated phases characterized by the development and dynamics of a pioneer and a mature forest species guild, respectively. Pioneer species dominated early recruitment (until ca. 10 years after abandonment), and declined before slower growing mature-forest species became dominant or reached maximum development rates (after 40–45 years). Pioneers promoted their replacement early in succession, while mature-forest species recruited and grew constantly throughout the process, with their lowest mortality coinciding with the peak of pioneer abundance. In contrast to prevailing stochastic views, we observed an orderly, community driven series of changes in this dry forest secondary succession. Chronosequences thus represent a valuable approach for revealing system-specific successional pathways, formulating hypotheses on causes and mechanisms and, in combination with repeated sampling, evaluating the effects of vegetation dynamics in pathway variation.     

Vegetation convergence during early succession on coal wastes: a 6‐year permanent plot study

Question: Does the course of succession on a coal mine restored by hydroseeding converge with a reference community in terms of species composition and vegetation structure? What is the rate of succession on restored areas? How does the balance between local colonization and extinction rates change during succession? Which species group (native or hydroseeded) determines the successional process? Location: Large reclaimed coal mine in the north of Palencia province, northern Spain (42°50′N, 4°38′W). Methods: Between 2004 and 2009 we monitored annually vascular plant species cover in nine permanent plots (20 m² each) at a restored mine; these plots were structured to account for site aspect (north, south and flat). Three identical permanent plots were established in the surrounding reference community and monitored in 2004 and 2009. We used detrended correspondence analysis to assess successional trends and rates of succession, generalized linear mixed models to derive patterns of vegetation structural changes and turnover through time, and Huisman–Olff–Fresco modelling to illustrate response of individual species through time. Results: The three restored mine areas exhibited a successional trend towards the reference community through time, although speed of convergence differed. However, after 6 years the restored sites had diverged considerably and this was greater than the dissimilarity reduction with respect to the reference community. Richness, diversity and native species cover increased linearly through time, whereas hydroseeded species cover decreased. Success of hydroseeded species initially differed in the three areas, and this was negatively related with native species colonization rates. Response patterns through time of ten hydroseeded and 20 most common native species are described. Conclusions: Vegetation structural parameters rapidly converged with the reference community, whereas compositional convergence needed much longer. At the same time, successional composition trajectories and rates were related to site properties (here aspect).     

Some factors affecting successional change on uranium mine wastes: Insights for ecological restoration

Question: Are growth form and dispersal-mode replacement during vegetation succession in semi-arid Mediterranean conditions affected by the starting quality of the substrate and by site aspect? Location: Central-western Spain. Methods: We monitored successions on three waste materials left after uranium mining: unbroken waste, broken waste and wastes amended with a sandy material (Arkoses); both north and south aspects were also studied on each substrate. Results: The substrate starting quality had the greatest influence on spontaneous succession, separating the poorer quality substrates (broken and unbroken wastes) from the better ones (Arkoses) and two reference communities (Topsoil and Dehesa). The importance of aspect was confirmed then within each substrate type. Most species with a short life span (mostly annuals and a few biennials), together with some woody species on Arkoses, showed no response to age (years following the deposition of new soil). Others short-lived species declined over time on the poorer wastes but not on the better Arkoses. There was a tendency for life form replacement (from thero-phytes to hemicryptophytes) during succession only on the poorer-quality substrates. No dispersal-mode replacement sequence was found. Conclusion: Improving the abiotic conditions of the substrate had a great effect on vegetation succession, but this effect was modified by aspect. Aspect took longer to induce differences in floristic composition on the poorer substrates, where succession was slower. Some trends in species responses to successional change were found by considering species traits, particularly life-form.     

Trends in monthly abundance and species richness of carabids over 33 years at the Kaiserstuhl,southwest Germany

Recent studies hint at consistent declines of insect abundance across taxa. However, detailed data from long-term surveys are rare in ecological studies, and yet are required in order to accurately infer trends and their causes. In the following, we analyse a dataset from pitfall traps sampled at a monthly resolution over a 33-year period (1979–2011) to investigate the activity density and species richness of ground beetles (Carabidae). The study site, an unmanaged xerophytic grassland amidst intensively managed vineyards in southwest Germany, underwent terracing for viticulture just before data collection began, which led the early community composition to be defined by succession. As this ecological situation initially conflates succession and other long-term trends, we define three successional phases (initial, early, and late), allowing us to separate trends over time. We use a generalised additive modelling approach to factor in the main drivers of the community trends in ground beetles, noticeably vegetation and climate. Both vegetation cover and climatic conditions had a small effect on carabid counts and species richness, and there was a notable decrease in both counts and species richness of 60% and 32%, respectively, during the late successional phase. An analysis of functional traits revealed an overall increase in community weighted mean biomass and a correlation of habitat preference with annual precipitation. Our results add to the growing volume of evidence pointing to a decline in insect populations both in central Europe and globally.     

Old-field successional dynamics on the Inland Pampa,Argentina

Abstract. The first 10 yr of old-field successional dynamics on the Argentine Inland Pampa were studied on a series of adjacent plots established consecutively between 1978 and 1989. We examined differences in species abundance patterns among plots in order to detect the spatial and temporal variability of succession. Perennial grasses steadily increased in cover and replaced the dominant annual species after 5 yr. Pioneer dicots persisted in older seral stages with 20 — 23 species/plot. Overall, exotic species (mostly the grasses Lolium multiflorum and Cynodon dactylon) contributed much to the plant cover in these communities. Native grasses comprised 45 % of total cover at years 7 — 10 of succession, but occurred with less than 7 species/plot. Substantial variation was found in the successional pathway, which reflected the particular sequence from annual forbs to short-lived and perennial grasses in the various plots. The course of succession was apparently influenced by a 2-yr period of unusually high rainfall. Deyeuxia viridiflavescens, a native perennial grass virtually absent before the wet period, spread over the study area and dominated seral communities for 3 yr, irrespective of plot age. Climatic conditions thus affected the successional turnover of life forms by increasing the rate of colonization by perennial grasses. We further point out the constraints imposed on secondary succession by the life histories of ‘available’ species.     

Successional trends in standing dead biomass in Mediterranean basin species

Question: Landscape models of fire occurrence in ecosystems assume that the time since the last fire determines vegetation flammability by enabling the accumulation of dead biomass. In this study we ask if Mediterranean basin shrublands respond to these models or, on the contrary, if initial successional stages in these ecosystems could be more flammable than later stages. Location: Mediterranean shrubland in the Valencia region, eastern Spain. Methods: Using different stages of vegetation development (5, 9, 14 and 26 years since the last fire), we first study the structural comiosition of the above‐ground biomass in 375 individuals of nine woody species. Then, we measure how the standing dead biomass varies during succession, taking into account the surface cover of each species and the quantity of total dead biomass accumulated in different successional stages (3, 9, 14 and 26 years since the last fire). Results: The largest amount of standing dead biomass at the plant community level is observed in the middle stages of the succession. Early successional species, such as Cistus spp., Ulex parviflorus and Pinus halepensis, have a higher percentage of standing dead biomass at earlier stages in the succession than species typical of later successional stages, e.g. Juniperus oxycedrus, Quercus coccifera and Quercus ilex. Conclusions: The results suggest that monotonic increase in fire hazard with increasing stand age is not necessarily the rule in Mediterranean basin shrublands, since early successional species may accumulate large amounts of standing dead biomass and thus promote fire at early successional stages.     

Changes in the structure and heterogeneity of vegetation and microsite environments with the chronosequence of primary succession on a glacier foreland in Ellesmere Island, high arctic Canada

Primary plant succession was investigated on a well-vegetated glacier foreland on Ellesmere Island in high arctic Canada. A field survey was carried out on four glacier moraines differing in time after deglaciation to assess vegetation development and microsite modification in the chronosequence of succession. The results showed evidence of directional succession without species replacement, which is atypical in the high arctic, reflecting the exceptionally long time vegetation development. During this successional process, Salix arctica dominated throughout all moraines. The population structures of S. arctica on these moraines implied the population growth of this species with progressing succession. The population density of S. arctica reflected the abundance of vascular plants, suggesting that development of the plant community might be related to structural changes and the growth of constituting populations. Through such growths of the population and the whole community with progressing succession, the spatial heterogeneity of vegetation gradually declines. Moreover, this vegetation homogenization is accompanied by changes in the spatial heterogeneity of microsite environments, suggesting significant plant effects on the modification of microsite environments. Accordingly, it was concluded that the directional primary succession observed on this glacier foreland is characterized by the initial sporadic colonization of plants, subsequent population growths, and the community assembly of vascular plants, accompanied by microsite modification.     

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.     

Development of vegetation patterns in early primary succession

Question: We investigated colonisation filters in early plant community development on a glacial outwash plain. We asked if these were related to seed limitation or to a lack of safe sites, if topographical heterogeneity affected species patchiness and how species life cycles influence successional trajectories. Location: An outwash plain (Skeiðarársandur) in southeast Iceland. Methods: We identified surface heterogeneity at two different scales, ca. 10–15 cm (larger stones and established plants) and ca. 50 m (shallow depressions representing dry river beds) at two study sites. We quantified species cover, flowering plant density, seed production, seed rain, seed bank density, seedling emergence and seedling survival from June 2005 to June 2007 for the whole plant community, and measured seed production for five species. Results: Mean vegetation cover was <2.5% at the sites. Low emergence rates and high seedling mortality were the two main recruitment filters. Only 1.4% of seedlings emerging in 2005 survived into the 2007 growing season. Topographical heterogeneity had little effect on plant colonisation. High annual variation was recorded, and the two study sites (ca. 2 km apart) differed in their colonisation success. Of the five species, establishment of Cerastium alpinum and Silene uniflora was most limited by lack of seeds, whereas establishment of Luzula spicata, Poa glauca and Rumex acetosella was most limited by safe sites. Conclusions: We conclude that colonisation processes and patterns in early primary succession on Skeiðarársandur were largely influenced by stochastic factors.     

Primary succession of Bistorta vivipara (L.) Delabre (Polygonaceae) root‐associated fungi mirrors plant succession in two glacial chronosequences

Glacier chronosequences are important sites for primary succession studies and have yielded well‐defined primary succession models for plants that identify environmental resistance as an important determinant of the successional trajectory. Whether plant‐associated fungal communities follow those same successional trajectories and also respond to environmental resistance is an open question. In this study, 454 amplicon pyrosequencing was used to compare the root‐associated fungal communities of the ectomycorrhizal (ECM) herb Bistorta vivipara along two primary succession gradients with different environmental resistance (alpine versus arctic) and different successional trajectories in the vascular plant communities (directional replacement versus directional non‐replacement). At both sites, the root‐associated fungal communities were dominated by ECM basidiomycetes and community composition shifted with increasing time since deglaciation. However, the fungal community's successional trajectory mirrored the pattern observed in the surrounding plant community at both sites: the alpine site displayed a directional‐replacement successional trajectory, and the arctic site displayed a directional‐non‐replacement successional trajectory. This suggests that, like in plant communities, environmental resistance is key in determining succession patterns in root‐associated fungi. The need for further replicated study, including in other host species, is emphasized.