Biogeographic synthesis of secondary succession rates in eastern North America

Aim Mechanistic models of old‐field plant succession have generally lacked a broader biogeographic context in which climate, soils and species pools could play a significant role in mediating succession rates. We examine broad‐scale patterns in old‐field succession and introduce a hierarchical conceptual model to incorporate potential mechanisms operating at multiple spatial scales. Location We reviewed secondary succession studies across the Eastern Deciduous Forest (EDF) of North America. Methods We collected data from all published studies that reported the time necessary for woody species to reach either 10% or 50% cover in old‐field systems throughout the EDF. We used regression approaches to determine whether rates of succession are controlled by climatic and edaphic factors that vary at broad spatial scales. Results We found that the rate at which woody species colonize and dominate old fields decreases significantly with latitude. Rates of woody succession were highly correlated with both annual temperature (growing degree‐days for years to 10% and 50% cover) and measures of soil fertility (cation exchange capacity and reported pre‐agroindustrial (1930) maize yields for years to 10% cover), all of which show a significant latitudinal trend across the EDF. Main conclusions We suggest that the driver of this pattern is geographic variation in the intensity of competition between herbaceous and woody species, which we relate to the effects of (1) temperature regime, (2) edaphic factors related to soil fertility, and (3) plant traits, which may vary latitudinally in response to climate and edaphic factors. Although insufficient data exist to evaluate the relative importance of these factors, we argue that research in this area is necessary to gain an understanding of how future landscapes will be affected by global climate and land use change.     

Interacting effects of herbivory and fertility on a synthesized plant community

1 Outdoor microcosms were used to investigate the effects of invertebrate herbivory on plant community composition, and thereby infer possible effects on the rate of secondary succession, at differing levels of soil fertility.
2 A mixture containing 24 grassland plant species of widely different functional types was established, with 12 microcosms at each of three fertility levels. Four generalist herbivores ( Helix aspersa , Cepaea hortensis , Arianta arbustorum and Sitobion avenae ) were added to half of the microcosms. Above-ground biomass of each species was harvested after 2 years. Reproductive variables were also measured for one species, Poa annua .
3 At both moderate and high soil fertility generalist invertebrate herbivores fed selectively on early successional, fast-growing species, thus increasing the relative abundance of later successional, slow-growing species. This supports the hypothesis that herbivory increases the rate of secondary succession. Flowering and viable seed production of early successional ephemerals was also reduced by the invertebrate herbivores across a wide range of soil fertility. This would seriously reduce the ability of a species to persist in the community, thereby further hastening the rate of succession.     

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.     

Early succession of butterfly and plant communities on set-aside fields

 Hypotheses on secondary succession of butterfly and plant communities were tested using naturally developed 1- to 4-year-old set-aside fields (n = 16), sown fields (n = 8) and old meadows (n = 4) in 1992 in South Germany. Pioneer successional fields (1st and 2nd year of succession, dominated by annuals) and early successional fields (3rd and 4th year of succession where perennials, especially grasses became dominant) had fewer plant species than mid-successional fields (old meadows). In contrast to established hypotheses, mean number of plant species decreased from 1- to 4-year-old set-aside fields. Species richness of butterfly communities did not change during the first four years of succession, but species composition changed greatly. Pioneer successional fields were characterized by (1) specialized butterflies depending on annual pioneer foodplants (e.g. Issoria lathonia), and (2) species preferring the pioneer successions despite their host plants being more abundant on early and mid-successional fields (e.g. Papilio machaon). The variability in butterfly species richness was best explained by flower abundance which was closely correlated with plant species richness. Species whose abundance was correlated with habitat connectivity were significantly smaller than species which correlated with flower abundance. Numbers of caterpillar species were correlated with numbers of adult butterfly species. Life-history features of butterflies changed significantly from pioneer to early and mid-successional fields. We found decreasing body size and migrational ability, decreasing numbers of species hibernating as imago, decreasing numbers of generations and increasing larval stage duration with age of succession, but, contrary to expectation, host plant specialization, numbers of egg-cluster laying species and egg diameter did not change with successional age. Received 18 September 1995 / Accepted: 17 July 1996     

Dynamics of vesicular-arbuscular mycorrhizae during old field succession

Summary The species composition of vesicular-arbuscular mycorrhizal (VAM) fungal communities changed during secondary succession of abandoned fields based on a field to forest chronosequence. Twenty-five VAM fungal species were identified. Seven species were clearly early successional and five species were clearly late successional. The total number of VAM fungal species did not increase with successional time, but diversity as measured by the Shannon-Wiener index tended to increase, primarily because the community became more even as a single species, Glomus aggregatum, became less dominant in the older sites. Diversity of the VAM fungal community was positively correlated with soil C and N. The density of VAM fungi, as measured by infectivity and total spore count, first increased with time since abandonment and then decreased in the late successional forest sites. Within 12 abandoned fields, VAM fungal density increased with increasing soil pH, H₂O soluble soil C, and root biomass, but was inversely related to extractable soil P and percent cover of non-host plant species. The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.     

Temporal patterns in rates of community change during succession

While ecological dogma holds that rates of community change decrease over the course of succession, this idea has yet to be tested systematically across a wide variety of successional sequences. Here, I review and define several measures of community change rates for species presence-absence data and test for temporal patterns therein using data acquired from 16 studies comprising 62 successional sequences. Community types include plant secondary and primary succession as well as succession of arthropods on defaunated mangrove islands and carcasses. Rates of species gain generally decline through time, whereas rates of species loss display no systematic temporal trends. As a result, percent community turnover generally declines while species richness increases--both in a decelerating manner. Although communities with relatively minor abiotic and dispersal limitations (e.g., plant secondary successional communities) exhibit rapidly declining rates of change, limitations arising from harsh abiotic conditions or spatial isolation of the community appear to substantially alter temporal patterns in rates of successional change.     

清除优势草本植物对西双版纳飞机草-马唐群落中木本幼苗的影响

在农业弃耕地上通常会出现由草本植物占优势的演替早期群落。为研究优势草本植物对群落中木本幼苗及群落微环境的影响以及外来入侵植物和本地植物对木本幼苗的影响有何差异,在西双版纳地区选取由外来入侵植物飞机草和本地植物马唐共同占优势的演替早期群落,实施物种清除实验,观测样地中木本幼苗的高增长及死亡补充情况,同时对群落微环境进行观测。结果显示,清除优势种显著提高了木本幼苗的高增长,降低了其死亡率,并且使新增幼苗数量有所提高。清除优势种对0-50cm幼苗高增长和死亡率的影响均大于50-100(或200)cm幼苗。清除处理显著提高了样地的冠层下可见天空比例,但对土壤含水量和土壤养分的影响并不显著。飞机草和马唐均能形成浓密的冠层,通过对光的竞争抑制木本幼苗的生长,且这两者的抑制作用没有显著差异。该地区次生林中常见的先锋树种在研究样地内均有幼苗存在,但其生长却受到优势草本植物的强烈抑制。因此,对演替早期群落中草本植物的控制与管理应同时注意外来入侵物种与本地杂草。     

Woody species colonisation in relation to habitat productivity

A study was conducted to analyse the effect of habitat productivity on woody species colonisation. Three soil types were distinguished: a relatively poor sandy soil type (1), a somewhat richer sandy type (2) and a relatively rich sandy loamy type (3). Chronosequences were established on these three soil types of 38 (type 1), 20 (type 2) and 54 years (type 3) after abandonment. In total 117 vegetation relevées were used to analyse life form change and species responses during old field succession via regression models. On the rich soil type the colonisation rate of woody species was slower than on the poor soil type. This can be explained by higher abundances of perennial species during the first 20 years after abandonment on the rich soil type in contrast to the poor soil type. Perennial species may delay the woody species colonisation. First they close the bare ground which inhibits germination and next they compete with woody seedlings for light, water and nutrients. The effect of habitat productivity on woody species colonisation can only be determined appropriately by taking life history traits into account. Early successional 'pioneer' woody species dispersed by wind have less difficulties colonising old fields than late successional 'forest' species; they colonise old fields prior to the development of a dense perennial sward. Forest species depend on animals to be dispersed which are attracted by vegetation structure. In ± 30 years on the poor soil type and in ± 45 years on the rich soil type woody species become dominant relative to other life forms. Forest species like Quercus robur L. invaded relatively early (<5 years) in contrast to other studies which probably coincides with the distance to seed sources (forest edges).     

Secondary succession, soil formation and development of a diverse community of oribatids and saprophagous soil macro-invertebrates

Investigations into different stages of secondary succession (from a wheat field to a beechwood on Threstone; Northern Germany) demonstrated the formation of a carbon rich top soil in later successional stages. Parallel to changes in plant species and soil formation, there were also changes in species composition and diversity of saprophagous macro-invertebrates (Lumbricidae, Diplopoda, Isopoda) and oribatid mites (Acari: Oribatida). Diversity of diplopod and isopod species increased after cessation of cultivation, but in a late successional stage (ca 50 y-old fallow, ash-dominated wood) species number of diplopods and isopods declined strongly. In comparison with the other soil invertebrate groups, species composition of earthworms among the sites was more similar. Accumulation of soil C was assumed to be related to wood formation and occurrence of woody debris and recalcitrant leaf litter of beech trees. Incorporation of recalcitrant litter materials by earthworm species living in the upper mineral soil presumably contributed significantly to accumulation of soil C. Accumulation of soil C was accompanied by the development of an oribatid mite community rich in species. In early successional stages oribatids predominantly colonized the litter layer, while most oribatid mites of the beechwood inhabited the upper mineral soil. Maximum diversity of oribatid mites in the beechwood is assumed to be related to instability of the mineral soil caused by earthworm activity. Changes in species composition and diversity are discussed considering succession theory. Even soil invertebrates of similar trophic groups appear to respond very differently to successional changes. It is concluded that conservation strategies to maintain high diversity of soil invertebrates are most likely to be successful if a wide range of habitats of different successional stages is included.     

云南松林次生演替阶段土壤细菌群落的变化

土壤细菌多样性是维持森林生态系统功能的关键因子,森林演替是影响其动态变化的重要因素。研究云南松林不同演替阶段土壤细菌群落结构及其多样性的变化规律,有助于深入理解森林生态系统恢复过程的驱动机制。本研究以云南省永仁县皆伐后形成的针叶林、针阔混交林和常绿阔叶林为对象,基于Illumina Hiseq高通量测序技术,分析森林演替过程中土壤细菌群落组成、结构、多样性及其影响因子的变化。结果表明: 土壤细菌的种群分类单元、Ace指数、Chao1指数和Shannon指数均随着演替进行呈减少趋势,演替早期阶段土壤的细菌总数、菌群丰富度及复杂程度最高。不同演替阶段细菌群落结构存在显著差异,其中,针阔混交林的差异最大,变形菌门和酸杆菌门为各演替序列共有的优势类群,放线菌门、绿弯菌门和Patescibacteria是演替早期的优势类群,且随着演替进行呈现减少趋势;变形菌门和WPS-2相对多度随演替进行呈增加趋势。土壤pH和乔木层物种丰富度是驱动次生演替过程中土壤细菌群落组成变化的关键因子。随着演替的进行,土壤细菌多样性减少,群落组成差异加大。     

Succession of bee communities on fallows

Wild bee communities were studied on one- to five-year-old set-aside fields with naturally developed vegetation (n = 20). and old orchard meadows (n = 4) to analyse effects of secondary succession on species diversity, resource use and associated life history traits. General theory predicts a steady increase of species richness with age of succession. In contrast, we found a first maximum in species richness of bees on two-year-old set-aside fields and a second on old meadows. Successional changes of bee communities were related to changes of vegetation. The transition from pioneer successional stages, dominated by annuals, to early successional stages, dominated by perennials, resulted in the highest species richness of flowering plants in the second year within the first five years of succession. Species richness of flowering plants was the best predictor variable for species richness of bees, whereas the cover of flowering plants correlated with the abundance of bees. Annual plants were visited more often and perennials less often than expected from their flower cover. Halictidae tended to prefer flowers of annuals, whereas Megachilidae. Apidae and Anthophoridae significantly preferred perennials. In departure from successional theory, body size, proportion of specialised bees and proportion of parasitic bees did not significantly increase with successional age, but number of generations and the proportion of soil-nesting bees decreased with successional age. Comparison of different management types showed that set-aside fields with naturally developed vegetation supported much more specialised and endangered bee species than set-aside fields sown with Phacelia tanacetifolia.     

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.     

Population structure and succession in temperate forests of southwestern Japan

In the temperate forests of southwestern Japan, the population density of woody plants in the community increases in the early stage of secondary succession, reaches a peak in the old oak-chestnut forest, and decreases towards the climax beech forest. The species richness and diversity of woody plants also show a trend similar to that of the population density. The canopy-tree population decreases in the course of the succession while the basal area increases, showing a self-thinning process. The species richness, diversity and population density of herbaceous plants are much influenced by the dominance of the bamboo, Sasa palmata. The life-history traits of trees, lower trees and shrubs are discussed in relation to their shoot system, reproductive pattern and successional processes.     

The role of plant resources in forest succession: changes in radiation, water and nutrient fluxes, and plant productivity over a 300-yr-long chronosequence in NW-Germany

In the past insufficient attention has been paid to quantitative measurements of resource fluxes in ecosystems that undergo successional change. In this study, simultaneous changes in seven plant resources (photosynthetically active radiation (PAR), water, nitrogen, phosphorus, calcium, magnesium and potassium) are quantified by a chronosequence approach for a 300-yr-long secondary succession on poor soil from Calluna vulgaris heathland to Fagus sylvatica-Quercus petraea late-successional forest (heathland-to-forest succession).Above-ground net primary production increases sevenfold, and total above-ground phytomass about fortyfold during heathland-to-forest succession. Plant organs that capture resources increase much more slowly (leaf area index: threefold; fine root biomass: 1.3-fold). The increase in productivity is based both on higher absorptivity and conversion efficiency of PAR by the canopies of the successional plants.Accumulation of organic material on the forest floor significantly improves soil water availability. Evapotranspiration losses increase early in succession as the growing vegetation increases in both height and leaf area but tend to decrease again in the late-successional community. Drainage losses are at their minimum at the conifer-dominated pioneer forest stage.Accumulation of available nutrients in the soil is a key process in heathland-to-forest succession that significantly improves plant nutrient availability but leads to only minor changes in carbon/nutrient ratios and humus quality. Litter decomposition rates increase and result in a more rapid nutrient turnover in late successional stages. External nutrient inputs (from the atmosphere and soil weathering) significantly contribute to plant nutrient supply early in succession, whereas the internal cycling of nutrients through litter fall and nutrient mineralisation by far exceeds external inputs at the late stages.Vitousek & Reiners' (1975) ecosystem nutrient loss hypothesis is supported by the heathland-to-forest succession data. Odum's (1969) hypotheses on how nutrient cycles change during the course of succession is, in one part, rejected, in part supported. Tilman's (1988) hypothesis on nutrient limitation early, and light limitation late in primary succession is rejected.     

Litterfall and Decomposition in Relation to Soil Carbon Pools Along a Secondary Forest Chronosequence in Puerto Rico

Secondary forests are becoming increasingly widespread in the tropics, but our understanding of how secondary succession affects carbon (C) cycling and C sequestration in these ecosystems is limited. We used a well-replicated 80-year pasture to forest successional chronosequence and primary forest in Puerto Rico to explore the relationships among litterfall, litter quality, decomposition, and soil C pools. Litterfall rates recovered rapidly during early secondary succession and averaged 10.5 (± 0.1 SE) Mg/ha/y among all sites over a 2-year period. Although forest plant community composition and plant life form dominance changed during succession, litter chemistry as evaluated by sequential C fractions and by ¹³C-nuclear magnetic resonance spectroscopy did not change significantly with forest age, nor did leaf decomposition rates. Root decomposition was slower than leaves and was fastest in the 60-year-old sites and slowest in the 10- and 30-year-old sites. Common litter and common site experiments suggested that site conditions were more important controls than litter quality in this chronosequence. Bulk soil C content was positively correlated with hydrophobic leaf compounds, suggesting that there is greater soil C accumulation if leaf litter contains more tannins and waxy compounds relative to more labile compounds. Our results suggest that most key C fluxes associated with litter production and decomposition re-establish rapidly—within a decade or two—during tropical secondary succession. Therefore, recovery of leaf litter C cycling processes after pasture use are faster than aboveground woody biomass and species accumulation, indicating that these young secondary forests have the potential to recover litter cycling functions and provide some of the same ecosystem services of primary forests.     

Spontaneous Establishment of Woody Plants in Central European Derelict Sites and their Potential for Reclamation

The performance of woody plants was analyzed in 15 successional seres starting at bare ground in central European manmade habitats. The total cover of woody species after 10 years of succession was significantly related neither to initial soil moisture nor to nitrogen (expressed using Ellenberg indicator values). But the comparison of seres indicates that establishment of woody plants was easier under moderate environmental conditions and retarded in extreme habitats (dry, nutrient-poor, or acid). The arrival of the first woody plants was delayed in dry sites. No significant differences were found between primary and secondary seres, either with respect to the total cover of woody plants reached after 10 years of succession or considering the time of their arrival. In total, 24 woody species (10 shrubs and 14 trees) appeared in the series investigated. Their successional performance (in terms of the number of seres in which the species occurred and maximum cover reached in any sere) was not related to species traits (life strategy, type of mycorrhizae, mode of dispersal, diaspore weight), except for the regeneration strategy, species with seasonal regeneration by seeds were capable of creating higher cover. Betula pendula (European birch) was the most successful species in spontaneous succession, especially on moist sites. Practical suggestions for the management of particular habitats (sites disturbed by mining, sites reclaimed after acid rain deforestation, urban sites, abandoned fields) are provided regarding the establishment of woody plants.     

Spatial Patchiness of Litter, Nutrients and Macroinvertebrates during Secondary Succession in a Tropical Montane Cloud Forest in Mexico

We test the hypothesis that secondary succession in Tropical Montane Cloud Forest (TMCF) in Mexico is accompanied by an increase in the spatial structuring of litter resources, soil nutrient concentrations and the soil macroinvertebrate community at a within-plot scale (5–25 m). This increased spatial structuring is expected because secondary succession in these forests is associated with an increase in the diversity of trees that dominate the canopy. If each tree species generates a particular soil environment under its canopy, then under a diverse tree community, soil properties will be spatially very heterogeneous. Tree censuses and grid sampling were performed in four successional stages of a secondary chronosequence of TMCF. Variography was used to analyse spatial patterns in continuous variables such as nutrient concentrations, while Spatial Analysis by Distance Indices (SADIE) was applied to determine patchiness in the distribution of soil macroinvertebrate taxa. Secondary succession was found to be accompanied by the predicted increase in the spatial structuring of litter resources and the macroinvertebrate community at the within-plot scale. Spatial patterns in the macroinvertebrate community only became evident for all taxa in the oldest forest (100 years old). Patches with low Ca and Mg concentrations in early successional soils were associated with patches where pine litter was most abundant while those with low P concentrations in late successional stages were associated with patches where oak litter was most abundant. Results suggest that anthropogenic disturbance aboveground promotes a more homogeneous resource environment in the surface soil, which compared to older forests, sustains a less diverse and less spatially structured macroinvertebrate community.     

Alternate pathways of succession on the Lake Erie Islands

In order to determine the effect of prior land use on secondary succession, woody plant species abundances were determined on the Lake Erie Islands in 21 sites 20–90 yr post-abandonment: 11 former vineyards, 8 former orchards, and 2 former quarries. Detrended Correspondence Analysis ordination was used to compare the relative abundances of woody plants in the 21 successional sites to each other and to 5 mature upland forests and 3 mature lowland forests. Vineyards 20–45 yr post-abandonment had closed canopies 2–4 m tall and were dominated by the same mix of species dominating mature forests at similar elevations. Abandoned orchards 20–45 yr post-abandonment supported species common in mature forest floras as well as many shade-intolerant, pioneer tree and shrub species not found in mature stands. These differences in successional patterns seem to be related to differences in the suite of colonization sites presented upon abandonment, with orchards being dominated by open, bare ground, and vineyards by shaded ground receiving ca. 20% of full sunlight. Newly abandoned vineyards had appreciable ‘advanced regeneration’ of saplings of shade-tolerant species 3–7 yr old growing among the grapes. While former quarries remained open, marginal habitats with sparse, open canopies dominated by species typical of rocky cliff edges, the community structure of former vineyards and orchards converged toward each other and toward the composition of mature forests at similar elevations. Thus patterns of secondary succession varied with prior land use and were determined by the suite of colonization sites available at the time of abandonment.     

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.     

Enhancement of late successional plants on ex-arable land by soil inoculations

Restoration of species-rich grasslands on ex-arable land can help the conservation of biodiversity but faces three big challenges: absence of target plant propagules, high residual soil fertility and restoration of soil communities. Seed additions and top soil removal can solve some of these constraints, but restoring beneficial biotic soil conditions remains a challenge. Here we test the hypotheses that inoculation of soil from late secondary succession grasslands in arable receptor soil enhances performance of late successional plants, especially after top soil removal but pending on the added dose. To test this we grew mixtures of late successional plants in arable top (organic) soil or in underlying mineral soil mixed with donor soil in small or large proportions. Donor soils were collected from different grasslands that had been under restoration for 5 to 41 years, or from semi-natural grassland that has not been used intensively. Donor soil addition, especially when collected from older restoration sites, increased plant community biomass without altering its evenness. In contrast, addition of soil from semi-natural grassland promoted plant community evenness, and hence its diversity, but reduced community biomass. Effects of donor soil additions were stronger in mineral than in organic soil and larger with bigger proportions added. The variation in plant community composition was explained best by the abundances of nematodes, ergosterol concentration and soil pH. We show that in controlled conditions inoculation of soil from secondary succession grassland into ex-arable land can strongly promote target plant species, and that the role of soil biota in promoting target plant species is greatest when added after top soil removal. Together our results point out that transplantation of later secondary succession soil can promote grassland restoration on ex-arable land.