Possible positive-feedback mechanisms: plants change abiotic soil parameters in wet calcareous dune slacks

In this paper the results are presented from a mesocosm study of the effects of typical dune slack plants on the soil solution nutrient contents. In dune slack succession, early successional species often show radial oxygen loss (ROL) whereas their successor species do not show ROL. ROL has impact on abiotic soil parameters and therefore, affect the competitiveness of both species. Mesocosms with Littorella uniflora and Carex nigra, used as respectively a ROL and a non-ROL species, showed remarkable differences in soil solution parameters. Special attention was given to nitrogen, as it is the limiting resource in dune slack succession. Mesocosms with L. uniflora showed a higher nitrate content in the soil than mesocosms with C. nigra and the control. Moreover, estimating the nitrogen balance, a significantly higher fraction of nitrogen was missing in L. uniflora (57%) than in C. nigra (5%). The enhanced nitrogen loss in mesocosms with L. uniflora could act as a positive-feedback mechanism for early successional stages that slows down the vegetation development in early stages of dune slack succession towards the more-productive later stages. The mechanism could even lead to alternative stable states in dune slack succession.     

Microbial community composition but not diversity changes along succession in arctic sand dunes

The generality of increasing diversity of fungi and bacteria across arctic sand dune succession was tested. Microbial communities were examined by high‐throughput sequencing of 16S rRNA genes (bacteria) and internal transcribed spacer (ITS) regions (fungi). We studied four microbial compartments (inside leaf, inside root, rhizosphere and bulk soil) and characterized microbes associated with a single plant species (Deschampsia flexuosa) across two sand dune successional stages (early and late). Bacterial richness increased across succession in bulk soil and leaf endosphere. In contrast, soil fungal richness remained constant while root endosphere fungal richness increased across succession. There was, however, no significant difference in Shannon diversity indices between early and late successional stage in any compartment. There was a significant difference in the composition of microbial communities between early and late successional stage in all compartments, although the major microbial OTUs were shared between early and late successional stage. Co‐occurrence network analysis revealed successional stage‐specific microbial groups. There were more co‐occurring modules in early successional stage than in late stage. Altogether, these results emphasize that succession strongly affects distribution of microbial species, but not microbial diversity in arctic sand dune ecosystem and that fungi and bacteria may not follow the same successional trajectories.     

Arbuscular mycorrhizal fungal communities change among three stages of primary sand dune succession but do not alter plant growth

Plant interactions with soil biota could have a significant impact on plant successional trajectory by benefiting plants in a particular successional stage over others. The influence of soil mutualists such as mycorrhizal fungi is thought to be an important feedback component, yet they have shown benefits to both early and late successional plants that could either retard or accelerate succession. Here we first determine if arbuscular mycorrhizal (AM) fungi differ among three stages of primary sand dune succession and then if they alter growth of plants from particular successional stages. We isolated AM fungal inoculum from early, intermediate or late stages of a primary dune succession and compared them using cloning and sequencing. We then grew eight plant species that dominate within each of these successional stages with each AM fungal inoculum. We measured fungal growth to assess potential AM functional differences and plant growth to determine if AM fungi positively or negatively affect plants. AM fungi isolated from early succession were more phylogenetically diverse relative to intermediate and late succession while late successional fungi consistently produced more soil hyphae and arbuscules. Despite these differences, inocula from different successional stages had similar effects on the growth of all plant species. Host plant biomass was not affected by mycorrhizal inoculation relative to un‐inoculated controls. Although mycorrhizal communities differ among primary dune successional stages and formed different fungal structures, these differences did not directly affect the growth of plants from different dune successional stages in our experiment and therefore may be less likely to directly contribute to plant succession in sand dunes.     

Effects of microbial mats on germination and seedling survival of typical dune slack species in the Netherlands

The role of microbial mats in wet dune slack succession is often discussed. We tested if presence of microbial mats may retard dune slack succession by lowering the germination and seeding survival of successor species. This hypothesis was tested on a set of typical dune slack species of the Frisian Islands in two climate chamber experiments. The species were separated into early-, intermediate- and late successional species. There were large differences in germination rates between species (2% – >200% compared to the reference), but within a species the high germination rates were mostly found on sand without a microbial mat. Only the germination of Agrostis stolonifera appeared to be stimulated by the presence of a well-developed microbial mat, they were even higher than in the reference. Seedling survival also did not show different responses between successional stages. Seedlings placed on top of a microbial mat showed for most species lower survival rates compared to seedlings that were planted or placed on top of the sand. Growth was the only measured variable that differed between successional groups. Species of the early- and intermediate successional stages grew significantly better if a microbial mat was present whereas late successional species were not stimulated. Early and intermediate successional species seem to be favored by the presence of a microbial mat. An explanation for this may be that they can profit from the enhanced nitrogen availability caused by N₂-fixation by cyanobacteria in the microbial mat. This revised version was published online in June 2006 with corrections to the Cover Date.     

Pattern analysis in successional communities – An approach for studying shifts in ecological interactions

Abstract. Many ecological studies have addressed issues of vegetation spatial patterns in attempts to understand the processes generating them. We investigated changes in ecological processes during succession via the analysis of shrubs’ spatial patterns in a system of linear sand dunes, an arid ecosystem located in the Negev Desert in Israel during three consecutive years. We hypothesized that spatial patterns change from clustered to regular as succession progresses due to changes in the relative importance of facilitation and competition in this environment. In this ecosystem communities of early successional stages are frequently disturbed by high rates of sand movement, whereas in later successional stages sand stability is high. We mapped in the field individual shrubs on high‐resolution aerial photographs, and converted the digital images to a GIS data set. Using Ripley's K‐function we analysed spatial patterns at three levels: the single‐species level, among species and at the individual level, in three communities characterizing different successional stages. In the early successional communities we found clustered spatial patterns, in comparison with stable habitats where spatial patterns tended to be regular. We argue that these shifts in spatial patterns are indicative of the assumption that in this sand‐dune system ecological interactions change from facilitation to competition as succession progresses. Further, we argue that these interactions operate in different spatial scales at the different successional stages, and that the study of these processes should be conducted at the spatial scales specific to each community.     

Aphid herbivory as a potential driver of primary succession in coastal dunes

Herbivory is a major factor affecting both the performance and the fitness of the species composing a plant community and, ultimately, conditioning its temporal and spatial dynamics. Coastal dunes are a typical example of primary succession where different biotic and abiotic factors determine plant species occurrence; however, the effect of insect herbivory herein has remained little explored. To address this matter, we combined an observational study along a successional gradient with a green-house experiment to determine the occurrence and the impact of plant–aphid interactions. We focused on the species Schizaphis rufula, a widespread and abundant aphid associated with dune grasses in early stages of primary succession in Europe. Firstly, we studied aphid infestation rates on the dune grass Ammophila arenaria along a succession gradient in three locations of the North Sea coast to address the relationship between plant community composition and aphid occurrence; secondly, we tested the effect of aphid herbivory on a set of dune species typical for the different stages of succession. We found that the degree of aphid infestation was inversely correlated with the degree of dune fixation. The results of the experiment showed that aphid multiplication was significantly higher and its effect more pronounced on two early successional grass species, i.e. A. arenaria and Leymus arenarius. Here aphid multiplication resulted in a severe decrease in plant biomass; in late successional grass species, there was limited multiplication and no effect on biomass. The results of the field survey and the green-house experiment indicate that aphids show a clear preference for plants from early successional stages and, moreover, they have a greater impact on these plant species. All this supports the hypothesis of aphid herbivory as a driving factor of primary succession in coastal dunes.     

Can the restoration of natural vegetation be accelerated on the Chinese Loess Plateau? A study of the response of the leaf carbon isotope ratio of dominant species to changing soil carbon and nitrogen levels

For the heavily degraded ecosystem on the Chinese Loess Plateau, it would be of great significance if vegetation restoration could be accelerated anthropogenically. However, one major concern is that if the late successional species were planted or sown in degraded habitats, would they still be competitive in terms of some critical plant traits associated with specific habitats? Water use efficiency (WUE) is a major plant trait shaping the pattern of species turnover in vegetation secondary succession on the Loess Plateau. We hypothesized that if late successional stage plants could still hold a competitive advantage in terms of WUE, the prospects for an acceleration of succession by sowing these species in newly abandoned fields would be good. We tested this hypothesis by comparing the leaf C isotope ratio (δ¹³C) value (a surrogate of WUE) of dominant species from different successional stages at given soil C and N levels. Results indicated that leaf δ¹³C of the two dominant species that co-dominated in the second and third stages were significantly more positive than that of the dominant species from the first stage regardless of changing soil C and N. Yet the dominant species from the climax stage is a C₄ grass assumed to have the highest WUE. In addition, increasing soil nutrition had no effects on leaf δ¹³C of two dominant species in the late successional stage, indicating that dominant species from the late successional stages could still have a competitive advantage in terms of WUE in soil C- and N-poor habitats. Therefore, from the perspective of plant WUE, there are great opportunities for ecosystem restoration by sowing both dominant species and other species that co-occur in late successional stages in newly abandoned fields, for the purpose of enhancing species diversity and optimising species composition.     

Factors controlling soil development in sand dunes: evidence from a coastal dune soil chronosequence

Aerial photographs, maps and optically stimulated luminescence dates were combined with existing soil data to construct high resolution chronosequences of soil development over 140 years at a temperate Atlantic UK dune system. Since soil formation had progressed for varying duration under different climate and nitrogen deposition regimes, it was possible to infer their relative influence on soil development compared with location-specific variables such as soil pH, slope and distance to the sea. Results suggest that soil development followed a sigmoid curve. Soil development was faster in wet than in dry dune habitats. In dry dunes, rates were greater than in the literature: they increased with increasing temperature and nitrogen deposition and decreased with increasing summer gales. The combination explained 62% of the variation. Co-correlation meant that effects of nitrogen deposition could not be differentiated from temperature. In wet dune habitats rates increased with temperature and decreased with gales. The combination explained only 23% of the variation; surprisingly, rainfall was not significant. Effects of location-specific variables were not significant in either habitat type. Nitrogen accumulation was faster in wet than dry dune habitats, averaging 43 kg N ha^?1 per year overall. Nitrogen accumulation greatly exceeded inputs from atmospheric deposition, suggesting rates of input for biological N fixation are 10–60 kg N ha^?1 per year. Recent climate and/or nitrogen deposition regimes may have accelerated soil development compared with past rates. These data suggest the importance of changing climate on soil development rates and highlight the contribution of biological N fixation in early successional systems.     

Generation of Spatial Patterns in Boreal Forest Landscapes

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

Plant hydraulics and photosynthesis of 34 woody species from different successional stages of subtropical forests

It is important to understand the ecophysiological characters of plants when exploring mechanisms underlying species substitution in the process of plant succession. In the present study, we selected 34 woody species from different stages of secondary succession in subtropical forests of southern China, and measured their hydraulic conductivity, gas exchange rates, leaf nutrients and drought‐tolerance traits such as xylem resistance to cavitation, turgor loss point and carbon isotope ratio. Principal component analysis revealed that early‐, mid‐ and late‐successional species were significantly separated along axis 1, which was strongly associated with hydraulic‐photosynthetic coordination. In contrast to species distributed in late‐successional forest, early‐successional species had the highest hydraulic conductivity, net photosynthetic rates, photosynthetic nitrogen and phosphorus use efficiencies, but had the lowest photosynthetic water‐use efficiency. However, changes of the measured drought‐tolerance traits of the 34 species along the succession did not demonstrate a clear trend – no significant correlations between these traits and plant successional stages were found. Moreover, the trade‐off between hydraulic efficiency and safety was not identified. Taken together, our results suggested that hydraulic efficiency and photosynthetic function, rather than drought tolerance, play an important role in species distributions along plant succession in subtropical forests.     

A stoichiometric model of early plant primary succession

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

Diversity and community biomass depend on dispersal and disturbance in microalgal communities

The evidence for species diversity effects on ecosystem functions is mainly based on studies not explicitly addressing local or regional processes regulating coexistence or the importance of community structure in terms of species evenness. In experimental communities of marine benthic microalgae, we altered the successional stages and thus the strength of local species interactions by manipulating rates of dispersal and disturbance. The treatments altered realized species richness, evenness and community biomass. For species richness, dispersal mattered only at high disturbance rates; when opening new space, dispersal led to maximized richness at intermediate dispersal rates. Evenness, in contrast, decreased with dispersal at low or no disturbance, i.e. at late successional stages. Community biomass showed a non-linear hump-shaped response to increasing dispersal at all disturbance levels. We found a positive correlation between richness and biomass at early succession, and a strong negative correlation between evenness and biomass at late succession. In early succession both community biomass and richness depend directly on dispersal from the regional pool, whereas the late successional pattern shows that if interactions allow the most productive species to become dominant, diverting resources from this species (i.e. higher evenness) reduces production. Our study emphasizes the difference in biodiversity–function relationships over time, as different mechanisms contribute to the regulation of richness and evenness in early and late successional stages.     

Alternative stable states in a wet calcareous dune slack in The Netherlands

Abstract. Evidence is presented for the occurrence of alternative stable states in a wet calcareous dune slack on the Frisian island of Texel, The Netherlands. An early pioneer stage (0.5 kgm^?2 total standing crop) and a more productive later successional stage (2.9 kg m^?2) occur side by side, with sharp boundaries between them. The pioneer vegetation has been recorded at the site for more than 62 yr. These features indicate the occurrence of a positive‐feedback mechanism that has led to alternative stable states. Analyses of ground and surface water composition, and decalcification depths, indicated that hydrologically the study site can be characterized as a flow‐through slack, with exfiltration of calcareous groundwater on one side and infiltration of surface water on the other side of the slack. These differences in hydrological conditions have led to distinct differences in environmental conditions within the dune slack. The occurrence of the two successional stages can, however, not be explained by differences in hydrological conditions since both stages occur side by side in the centre of the dune slack. It is, therefore, more likely that biotic interactions are the cause of the vegetation pattern. Three possible mechanisms for feedback processes are discussed: (1) enhanced nitrogen loss; (2) sulfide toxicity and (3) nutrient accumulation in internal cycle.     

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.     

Steady and dynamic photosynthetic responses of seedlings from contrasting successional groups under low‐light growth conditions

To test the hypothesis that leaf‐level photosynthetic‐related traits might confer late successionals a competitive advantage over early successionals in low‐light growth conditions, steady photosynthetic assimilation‐ and dynamic photosynthetic induction‐related traits were examined in low‐light‐grown seedlings with contrasting successional status. Compared with the early successionals, late successionals as a group significantly exhibited lower leaf gas exchange rates. While late successionals required a longer time to respond to simulated sunflecks, they had lower rates of induction losses after sunflecks. Such photosynthetic induction traits allowed late successionals to more effectively utilize subsequent sunflecks. It was observed that plants with lower gas exchange rates responded more slowly to simulated sunfelcks, but they had lower rates of induction losses after sunflecks. In addition, the rate of response to sunflecks was positively correlated with the rate of induction loss after sunflecks across the successional status of species. A principal components analysis (PCA) demonstrated that early and late successionals were separated along the first axis of the PCA, and that early successionals were grouped on the right and were associated with higher gas exchange rates, fast responses to sunflecks, and rapid rates of induction loss after sunflecks; late successionals held an opposite pattern. Overall, our results suggest that smaller respiratory carbon losses and lower metabolic costs give late successionals a competitive advantage in low‐light growth conditions, that late successionals have an advantage over early successionals in utilizing sunflecks, and thus that the successional status of species are mainly associated with the leaf‐level photosynthetic‐related traits.     

Nitrate reduction capacity is limited by belowground plant recovery in a 32‐year‐old created salt marsh

Human activities have decreased global salt marsh surface area with a subsequent loss in the ecosystem functions they provide. The creation of marshes in terrestrial systems has been used to mitigate this loss in marsh cover. Although these constructed marshes may rapidly recover ecosystem structure, biogeochemical processes may be slow to recover. We compared denitrification and dissimilatory nitrate reduction to ammonium (DNRA) rates between a 32‐year‐old excavation‐created salt marsh (CON‐2) and a nearby natural reference salt marsh (NAT) to assess the recovery of ecosystem function. These process rates were measured at 5 cm increments to a depth of 25 cm to assess how plant rooting depth and organic matter accumulation impact N‐cycling. We found that, for both marshes, denitrification and DNRA declined with depth with the highest rates occurring in the top 10 cm. In both systems, N‐retention by DNRA accounted for upwards of 75% of nitrate reduction, but denitrification and DNRA rates were nearly 2× and 3× higher in NAT than CON‐2, respectively. Organic matter was 6× lower in CON‐2, likely due to limited plant belowground biomass production. However, there was no response to glucose additions, suggesting that the microbial functional community, not substrate limitation, limited nitrate reduction recovery. Response ratios showed that denitrification in CON‐2 recovered in surficial sediments where belowground biomass was highest, even though biomass recovery was minimal. This indicates that although recovery of ecosystem function was constrained, it occurred on a faster trajectory than that of ecosystem structure.     

Nutrient limitation and vegetation changes in a coastal dune slack

Abstract. Basiphilous pioneer species are among the most endangered plant species in The Netherlands. They find most of their refuges in young coastal dune slacks, especially on the Wadden Sea islands. For the purpose of nature management it is important to know which processes control the presence of basiphilous pioneer communities, and to learn about the nature of slacks harbouring the concerning successional sequences. In a large dune slack on the Island of Terschelling, we assessed soil nutrient status and tested for nutrient limitation in four chronosequential stages: 2, 6, 37 and ca. 80 yr of age. Stage 2 harboured a basiphilous pioneer vegetation; in the stages 3 and 4 a dense vegetation of dwarf shrubs and grasses occurred. Soil organic matter and nutrient concentrations in each stage were measured in 1991. In 1992 and 1993 fertilizers were applied to all stages to detect nutrient limitation. Rates of accumulation of organic matter, nutrients and above-ground biomass were estimated. When interpreted as successional stages, the different stages represent a sequence as expected on the basis of general successional theory. There was a peak in yearly nutrient accumulation between the 6- and 37-yr old stage and a steady state after ca. 80 yr. Between the first two and the latter two stages a shift occurred from allogenic to autogenic succession which correlated with a shift in emphasis from available nutrients to light availability as limiting resources. Basiphilous pioneer species suffered only deficiency of nitrogen, probably because of their low phosphorus requirements. It is concluded that in dune slack habitats, in addition to a low nutrient availability in general, a very low phosphorus availability favours basiphilous pioneer species to species showing co-limitation of nitrogen and phosphorus as found in some grasses and dwarf shrubs. A comparison between the effects of lime addition and the effects of nitrogen and phosphorus additions suggests that, in the early stages, soil buffering increases the availability of nitrogen and inhibits the availability of phosphorus. Sod cutting is an effective technique for restoring basiphilous pioneer vegetation, when slacks are acidified only superficially and buffering-mechanisms can be reactivated. Yearly mowing and removing of standing crop may prolong the life-span of basiphilous pioneer vegetation, when soil acidification has not yet dropped below pH 6.     

Interactions between mycorrhizal colonization and plant life forms along the successional gradient of coastal sand dunes in the eastern Mediterranean, Turkey

The mycorrhizal status of dune plant species in relation to their plant life forms was surveyed along a successional gradient of sand dune on the southern Mediterranean coast of Turkey. Roots of 64 dune plant species belonging to 30 families were collected from sand dune communities at four different successional stages: embryonic dunes (ED), mobile dunes (MD), fixed dunes (FD), and remnant dunes (RD). Of the plant species surveyed in all successional stages, 54 (84%) had formed mycorrhizal associations. Nonmycorrhizal plants with cryptophyte life forms predominated in the earlier successional stages (ED and MD), whereas the number and percent coverage of mycorrhizal plant species belonging to hemicryptophytes, phanerophytes, and chamaephytes generally increased with the stabilization of sand dunes. Arbuscular mycorrhizal (AM) colonization was found to be the dominant mycorrhizal type in ED, MD, and RD. But phanerophytes with dual colonization, AM and ectomycorrhizal, became the dominant life form with high plant coverage in the FD stage. Total percentage of mycorrhizal root length colonization showed significant positive correlations relating to soil parameters such as organic matter and nitrogen content, while negatively correlating to high soil reaction (pH).     

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

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