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.     

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.     

The influence of aspect on the early growth dynamics of hydroseeded species in coal reclamation areas

Question: Does aspect affect hydroseeding success and the development of vegetation during early vegetation establishment on the steep slopes of coal wastes during the reclamation process? Location: Open‐pit coal mine near Villanueva de la Peña, northern Spain. Methods: In the first year after hydroseeding, we monitored the dynamics of hydroseeded species in three permanent plots of 20 m² on north‐ and south‐facing slopes every two months. Soil properties and weather conditions were also monitored. Results: Aspect was related to total plant cover during early revegetation, and south‐facing slopes had the lowest cover. Aspect also influenced the early dynamics of hydroseeded grasses and legumes establishing on these slopes. Grass cover was greater on the north slope throughout the study, but differences in plant cover between north and south slopes appeared later for the legumes. Aspect also affected the relative contribution of both of grasses and legumes to the total plant cover, with grasses dominant on both northern and southern slopes, except during the summer on the southern slope. The species with the greatest difference in cover between the north‐ and south‐facing slopes were Festuca spp., Lolium perenne and Trifolium repens. Conclusion: In coal mine reclamation areas of Mediterranean climates, differences in the development of hydroseeded species depended on the slope of the coal mine reclamation areas, and this information is of importance to managers in selecting species for use in reclamation.     

Properties of native plant communities do not determine exotic success during early forest succession

Considerable research has been devoted to understanding how plant invasions are influenced by properties of the native community and to the traits of exotic species that contribute to successful invasion. Studies of invasibility are common in successionally stable grasslands, but rare in recently disturbed or seral forests. We used 16 yr of species richness and abundance data from 1 m² plots in a clearcut and burned forest in the Cascade Range of western Oregon to address the following questions: 1) is invasion success correlated with properties of the native community? Are correlations stronger among pools of functionally similar taxa (i.e. exotic and native annuals)? Do these relationships change over successional time? 2) Does exotic abundance increase with removal of potentially dominant native species? 3) Do the population dynamics of exotic and native species differ, suggesting that exotics are more successful colonists? Exotics were primarily annual and biennial species. Regardless of the measure of success (richness, cover, biomass, or density) or successional stage, most correlations between exotics and natives were non‐significant. Exotic and native annuals showed positive correlations during mid‐succession, but these were attributed to shared associations with bare ground rather than to direct biotic interactions. At peak abundance, neither cover nor density of exotics differed between controls and plots from which native, mid‐successional dominants were removed. Tests comparing nine measures of population performance (representing the pace, magnitude, and duration of population growth) revealed no significant differences between native and exotic species. In this early successional system, local richness and abundance of exotics are not explained by properties of the native community, by the presence of dominant native species, or by superior colonizing ability among exotics species. Instead natives and exotics exhibit individualistic patterns of increase and decline suggesting similar sets of life‐history traits leading to similar successional roles.     

The importance of topography and climate on short-term revegetation of coal wastes in Spain

The initial colonization phase is the crucial start point for succession and therefore for restoration. However, little is known about abiotic factors that influence the early stages of revegetation dynamics on restored coal mines, particularly in a Mediterranean climate. This information is crucial for improving our ability to reclaim land despoiled by mining. Here, we characterized the short-term plant community development in the first 3 years after hydroseeding on a topographically diverse reclaimed open-pit coal mines in Spain. Topography influenced both community composition and diversity producing different trajectories between the three different aspects (north-facing, south-facing, flat). Hydroseeded species provided most of the initial vegetation cover, which brought about most of the compositional differences between these aspects. We also found that the changing climate through the growing season also influenced floristic composition and diversity. Summer drought reduced the cover of hydroseeded perennial species and allowed native species to colonize. These results emphasize the need to take topography into account when developing reclamation management plans in Mediterranean ecosystems, and an acknowledgement that climate might drive the succession in the desired direction, hence increasing success in restoring mining impacts.     

Colonization and early succession on anthropogenic soils

Abstract. This paper reports on vegetation development on permanent experimental plots during five years of succession. Nine (1 m²) plots were filled with three typical substrates from man-made habitats of urban and industrial areas in the region of Berlin. The three substrates (a commercial ‘topsoil’, a ruderal ‘landfill’ soil and a sandy soil), differ in organic matter and nutrient contents. Relevés of species composition and percent cover of each species present were made monthly during the growing season from the start of vegetation development. This paper describes the different successional pathways on topsoil and ruderal soil and the colonization process on sandy soil. On topsoil, ruderal annuals are dominant in the first year and are replaced by short-lived perennials from the second year. Those species were replaced by long-lived perennial herbs (Ballota nigra or Urtica dioica) from the third year of succession onwards. On the ruderal land-fill soil the early successional stages are less sharp and the perennial Solidago canadensis is able to dominate within one year after the succession was initiated. On sandy soil there is still an ongoing colonization process, where pioneer tree species like Betula pendula and Populus nigra play a main role. The importance of ‘initial floristic composition’, the role of substrate for community structure and the peculiarities of successional sequences on anthropogenic soils in the context of primary and secondary successions are discussed.     

Plant colonization windows in a mesic old field succession

Abstract. Closed canopy vegetation often prevents the colonization of plant species. Therefore the majority of plant species are expected to appear at the initial phase of post‐agricultural succession in mesic forest environment with moderate levels of resources. This hypothesis was tested with data from the Buell‐Small Successional Study, NJ, USA, one of the longest continuous fine‐scale studies of old‐field succession. The study started in 1958, including old fields with different agricultural histories, landscape contexts, and times of abandonment. In each year of the study, the cover values of plant species were recorded in 48 permanent plots of 1 m² in each field. We analysed the temporal patterns of colonization at plot scale and related these to precipitation data and other community characteristics. The number of colonizing species decreased significantly after ca. 5 yr, coinciding with the development of a continuous canopy of perennial species. However, species turnover remained high throughout the whole successional sequence. The most remarkable phenomenon is the high inter‐annual variation of all studied characteristics. We found considerable temporal collapses of vegetation cover that were synchronized among fields despite their different developmental stages and distinctive species compositions. Declines of total cover were correlated with drought events. These events were associated with peaks of local species extinctions and were followed by increased colonization rates. The transitions of major successional stages were often connected to these events. We suggest that plant colonization windows opened by extreme weather events during succession offer optimum periods for intervention in restoration practice.     

Dynamic and static views of succession: Testing the descriptive power of the chronosequence approach

Chronosequence and permanent plot studies are the two most common methods for evaluating successional dynamics in plant communities. We combined these two approaches by re-sampling an old-field chronosequence at Cedar Creek Natural History Area (Minnesota, USA) to: (1) measure rates of secondary succession; and (2) to test the ability of the chronosequence approach to predict actual successional dynamics over a 14-year survey interval. For each of 19 chronosequence fields we calculated four complimentary indices of succession rate for community changes that actually occurred within each of these fields between 1983 and 1997. We found that measures of compositional dissimilarity, species turnover, and the change rates of perennial and native species cover over this 14-year period were all negatively correlated with field age, indicating that the rate of successional change in these old-fields generally declines over time. We also found that data collected from the initial static chronosequence survey (1983) accurately predicted many of the observed changes in species abundance that occurred between 1983 and 1997, but was a poor predictor of changes in species richness. In general, chronosequence re-sampling confirmed the validity of using the chronosequence approach to infer basic patterns of successional change.     

Patterns of early succession on bare peat in a Swiss mire after a bog burst

Questions: How does plant diversity (species richness, species abundance and rate of change) evolve in early succession on bare peat? Does succession converge towards one equilibrium stage or end up in several stages? Is there a regular pattern in succession velocity? Location: A mire in the calcareous Jura Mountains of northwest Switzerland. Method: Twenty‐one 1‐m² permanent plots on bare peat were used to monitor temporal stages over a 21‐year period (1988 to 2008) in a Swiss mire where a slide occurred in 1987. Species diversity and life forms were analysed based on Shannon's equitability index and cover. We used classification and metric ordination techniques to investigate patterns of successional rates and trends. The high temporal resolution of the survey allowed the pattern of succession velocity to be analysed. Results: Species richness increased continuously over the 21 years of succession. The highest cover throughout the study period was the life form sedge. Time trajectories of the 21 plots revealed three alternative pathways towards intermediate equilibrium stages in the first years, still not converging in the later stages. Changes in succession velocity reached a first maximum about 6 years after the slide had occurred and a second maximum 12 years later.     

Effects of disturbance intensity and frequency on early old‐field succession

Abstract. Early old‐field succession provides a model system for examining vegetation response to disturbance frequency and intensity within a manageable time scale. Disturbance frequency and intensity can interact with colonization and competition to influence relative abundance of earlier and later successional species and determine, respectively, how often and how far succession can be reset. We tested the joint effects of disturbance frequency and intensity on vegetation response (species richness, abundance, canopy structure) during the first six years of succession by clipping the dominant species (D) or all species (T) in spring and fall of each year (S), once per year in summer (Y1), each two years in summer (Y2), or each four years in summer (Y4). Vegetation response reflected disturbance effects on expansion of a later monospecific dominant perennial herb, Solidago altissima, and persistence of the early, richer flora of annuals. A more abundant and taller top Solidago canopy developed on plots clipped each 2 yr or less frequently. Plots clipped yearly or seasonally were richer, but had less abundant, shorter, and differently stratified canopy. Disturbance mediated the relative abundance of early and later successional species; however, frequency and intensity effects were not completely congruent. Persistence of a richer early successional flora increased through the most frequent disturbance (S), and was magnified by disturbance intensity. Disturbance as extreme as clipping all vegetation twice yearly did not cause a drop in species richness, but maintained the early successional community over the first six years of succession. We conclude that clipping disturbance influenced the rate of succession, but the early community could rebound through the range of disturbance frequency and intensity tested.     

Convergence of community composition during secondary succession on Zokor rodent mounds on the Tibetan Plateau

Aims The community succession theory is much debated in ecology. We studied succession on Zokor rodent mounds on the Tibetan Plateau to address several fundamental questions, among them: (i) During secondary succession, does the community composition converge towards one community state or multiple states depending on the initial colonization? (ii) Do mound communities located in different background communities exhibit different assembly trajectories?Methods In a sub-alpine meadow, we investigated a total of 80 mound communities at several successional stages in three different background communities resulting from different management histories and compared their changes in species composition. The distribution of plant communities over time was analyzed with quantitative classification and ordination methods. The co-occurrence patterns of species were evaluated at each successional stage, and the degree of convergence/divergence among communities was obtained by calculating two beta-diversity indices.Important findings During secondary succession, species richness of mound communities changed over time, and this change was dependent on the background community. Five life-form groups exhibited different dynamic patterns in species richness and plant cover. Community composition and the degree of species co-occurrence between communities increased over time since disturbance. There was much variation in species composition at earlier stages of succession, but communities on older mounds became more similar to each other and to their surrounding vegetation over the course of secondary succession. Post-disturbance succession of Zokor mound communities transitioned from 'multiple alternative states' to 'background-based deterministic community assembly' over time. Tradeoffs between competition and colonization, as well as the characteristics of different life-forms and mass effects within a limited species pool are the mechanisms responsible for convergence of mound communities.     

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.     

Effects of grassland succession on communities of small mammals in Illinois, USA

Native grasslands are among the most imperiled of the North American ecosystems, with only ∼4% of their pre-settlement area remaining, but some grassland habitats are being restored and maintained through such methods as prescribed burning and mowing, which may provide habitat for animal species endemic to this ecosystem. I determined how succession of the plant community, due to a four-year rotational burn in 16 grassland fragments, influenced species richness and local abundances of small mammals in Illinois, USA. Species richness was relatively low in grasslands that were recently burned and highest in older successional grasslands. The most abundant species, Microtus ochrogaster, M. pennslyvanicus, Peromyscus maniculatus, P. leucopus, and Reithrodontomys megalotis showed very different responses to succession; Microtus spp. were most abundant in older successional grasslands, preferring areas with more cover of bunchgrasses, whereas the other three species were relatively abundant in grasslands of all successional ages. P. maniculatus was most abundant in any habitat that had ample open ground. The grasslands at my study site are a mixture of restored and non-restored grasslands. Overall, adding additional time between burns and restoring more of the grasslands by planting bunchgrasses that are native to this area may increase abundances of most mammal species at my study site.     

Effects of Sewage Sludge on Plant Community Composition in Restored Limestone Quarries

The effects of sewage sludge, used to improve fertility of replaced soil, on vegetation were studied in limestone quarry restoration. Plant community growing in the first stages after sludge application was surveyed in six quarries of NE Spain. Areas with a mixture of sewage sludge and residual soil were compared to areas where the application consisted only of residual soil (a mixture of previous top soil and mine spoils). Sewage sludge was hypothesized to increase total biomass and cover, modifying species composition and delaying the early successional recover of the community. The results showed that both biomass and plant cover increased because of sewage sludge addition. The floristic composition was dominated by ruderal species that did not show any dependence on sewage sludge application. Convergence on similarity between sludge and control plots was not detected along a 5‐year period. Although species richness was significantly lower in sludge plots, diversity and equitability indexes did not show differences between treatments. The results did not show differences in the proportion of non‐native species. The proportion of legumes was lower in sludge plots. These results show that the plant communities resulting from the addition of sewage sludge to the soil used in limestone quarry restoration have more biomass and cover, but less number of species, and they do not show a clear trend to converge to those areas restored only with non‐amended soil.     

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’.     

Directionality,convergence, and rate of change during early succession in the Inland Pampa,Argentina

Abstract. We tested the following hypotheses forthe first five years of a grassland succession: (a) community changes are mainly directional and related to time after disturbance ratherthanto environmental fluctuations; (b) rates of succession decrease over time, and (c) plant communities in different plots converge on a similar composition within five years of succession. We tested those hypotheses using a Principal Components Analysis applied to data from four successional plots established in successive y ears in a large cropland in the Inland Pampa, Argentina. Community changes were correlated to the age of the plots, and unrelated to rainfall variability, a major environmental variable in grasslands. Successional rates were constant over the five years, probably because of the continued dominance of different annuals; we conclude that successional rates depend on the life history of the dominant species rather than on any emergent community property. We found no evidence of convergence ordivergence; we concluded that the results of successional studies may depend on the temporal and spatial scale of observation.     

Experimental analysis of patch dynamics and community heterogeneity in tallgrass prairie

Effects of fire and small-scale soil disturbances on species richness, community heterogeneity, and microsuccession were investigated in a central Oklahoma tallgrass prairie. In the fall of 1985, 0.2 m² soil disturbances were created on burned and unburned tallgrass prairie. Vegetation on and off disturbances was sampled at monthly intervals over two growing seasons. During the first growing season, the cover of forbs and annuals, and species richness were significantly greater on versus off disturbances, but these differences did not persist through the second year. The variation in species composition among disturbed plots (heterogeneity) was significantly greater compared to undisturbed areas throughout the study. Fire had no consistent effect on richness and heterogeneity of vegetation on soil disturbances but fire reduced heterogeneity on undisturbed vegetation. Rate of succession, based on an increase in cumulative cover of perennial grasses over time, did not differ among treatments during the first growing season. During the second year, rate of succession was significantly greater on burned soil disturbances compared to unburned soil disturbances. These results suggest that while small-scale soil disturbances have primarily short-lived effects on grassland community structure, disturbances do help to maintain spatial and temporal variation in tallgrass prairie communities. Unlike in undisturbed vegetation, however, species richness and heterogeneity on soil disturbances were little effected by fire, but the rate of colonization onto disturbances appeared to be enhanced by fire.     

Limits to convergence of vegetation during early primary succession

Questions: Primary succession, measured by changes in species composition, is slow, usually forcing a chronose‐quence approach. A unique data set is used to explore spatial and temporal changes in vegetation structure after a 1980 volcanic eruption. On the basis of data from a transect of 20 permanent plots with an altitudinal range of 250 m sampled through 2005, two questions are asked: Do changes along the transect recapitulate succession? Do plots converge to similar composition over time? Location: A ridge between 1218 and 1468 m on Mount St. Helens, Washington, USA. Methods: Repeat sampling of plots for species cover along a 1‐km transect. Floristic changes were characterized by techniques including DCA, clustering and similarity. Results: Species richness and cover increased with time at rates that decreased with increasing elevation. The establishment of Lupinus lepidus accelerated the rate of succession and may control its trajectory. Diversity (H) at first increased with richness, then declined as dominance hierarchies developed. Primary succession was characterized by overlapping phases of species assembly (richness), vegetation maturation (diversity peaks, cover expands) and inhibition (diversity declines). Each plot passed through several community classes, but by 2005, only four classes persisted. Succession trajectories (measured by DCA) became shorter with elevation. Similarity between groups of plots defined by their classification in 2005 did not increase with time. Similarity within plot groups converged slightly at the lower elevations. Despite similarities between temporal and spatial trends in composition, trajectories of higher plots do not recapitulate those of lower plots, apparently because Lupinus was not an early colonist. Any vegetation convergence has been limited to plots that are in close proximity.     

Long‐term effectiveness of sowing high and low diversity seed mixtures to enhance plant community development on ex‐arable fields

Questions: How is succession on ex‐arable land affected by sowing high and low diversity mixtures of grassland species as compared to natural succession? How long do effects persist? Location: Experimental plots installed in the Czech Republic, The Netherlands, Spain, Sweden and the United Kingdom. Methods: The experiment was established on ex‐arable land, with five blocks, each containing three 10 m × 10 m experimental plots: natural colonization, a low‐ (four species) and high‐diversity (15 species) seed mixture. Species composition and biomass was followed for eight years. Results: The sown plants considerably affected the whole successional pathway and the effects persisted during the whole eight year period. Whilst the proportion of sown species (characterized by their cover) increased during the study period, the number of sown species started to decrease from the third season onwards. Sowing caused suppression of natural colonizing species, and the sown plots had more biomass. These effects were on average larger in the high diversity mixtures. However, the low diversity replicate sown with the mixture that produced the largest biomass or largest suppression of natural colonizers fell within the range recorded at the five replicates of the high diversity plots. The natural colonization plots usually had the highest total species richness and lowest productivity at the end of the observation period. Conclusions: The effect of sowing demonstrated dispersal limitation as a factor controlling the rate of early secondary succession. Diversity was important primarily for its‘insurance effect’: the high diversity mixtures were always able to compensate for the failure of some species.     

Plant succession in areas of scorched and blown-down forest after the 1980 eruption of Mount St. Helens,Washington

Abstract. Patterns of plant succession were studied in areas of scorched and blown-down forest resulting from the 1980 eruption of Mount St. Helens, Washington. Changes in species abundance were observed for 7 years in permanent sample plots representing four post-disturbance habitats, or site types. Total plant cover and species richness increased with time on all site types. In blown-down forests supporting snowpack at the time of eruption, understory recovery was dominated by the vegetative regeneration of species persisting through disturbance. In forests without snowpacks, plant survival was poorer. Increases in cover and diversity were dominated first by introduced grasses, then by colonizing forbs characteristic of early successional sites. Epilo-bium angustifolium and Anaphalis margaritacea showed widespread recruitment and clonal expansion throughout the devastated area. As a result, species composition on previously forested sites converged toward that on formerly clearcut sites, where early serai forbs resprouted vigorously from beneath the tephra. Total plant cover and species diversity were poorly correlated with post-disturbance habitat and general site characteristics (e.g. distance from the crater, elevation, slope, and aspect). However, distributions of several life-forms (e.g. low sub-shrubs and tall shrubs) were strongly correlated with depth of burial by tephra and with cover of tree rootwads. Thus, early community recovery may reflect microsite variation or chance survival and recruitment rather than broad-scale gradients in environment or disturbance. Recovery of pre-disturbance composition and structure will undoubtedly be much slower than after other types of catastrophic disturbance. The rate and direction of community recovery will largely depend on the degree to which original understory species survived the eruption.