Long-term Succession Research Articles

Long-term succession of aquatic plants reconstructed from palynological records in a shallow freshwater lake

Aquatic plants in shallow freshwater lakes play a key role in stabilizing ecological function and providing valuable ecosystem services, yet they are severely degraded worldwide. An improved understanding of long-term aquatic plant succession is critical to investigate the potential driving mechanisms and to facilitate ecological restoration. In this paper, we reconstructed changes in the aquatic plant community over the past century based on palynological records from Changdang Lake, Middle and Lower Yangtze River Basin (MLYB), China. Our results reveal that aquatic plants in Changdang Lake have undergone three clear phases: emergent macrophytes dominated the aquatic vegetation in the 1900s–1970s, submerged macrophytes in the 1970s–1990s, and floating macrophytes increasingly after the 1990s. Significant changes in the aquatic plant communities were caused by increasing anthropogenic pressures, such as damming and nutrient loading from agriculture, aquaculture, and urbanization after the Chinese economic reform. We argue that Changdang Lake is currently in a transition phase between a macrophyte-dominated state and an algae-dominated state. Our palynological record is different from many contemporary studies, which suggest submerged plants dominated most lakes in this region before the 1950s. We suggest that the return of the aquatic plants to their 1970s–1980s state would be a realistic target for lake restoration. Our results show that palynological records can reveal long-term dynamics of macrophytes in shallow lakes for sustainable lake restoration and management.

Dynamics of understorey biomass, production and turnover associated with long-term overstorey succession in boreal forest of Canada

Understorey vegetation hosts the most species diversity in temperate and boreal forests and contributes substantially to ecosystem functions. Despite its ecological importance, the dynamics of understorey biomass, production and turnover rates following stand-replacing disturbance and throughout forest succession remain poorly understood. Using a replicated chronosequence spanning 8, 16, 34, 98, 146 and 210 years following fire, we examined the dynamics of understorey biomass, production and turnover associated with stand development and overstorey types (broadleaf, mixedwood, and conifer) in the central boreal forest of Canada. Across all ages and overstorey types, biomass, production and turnover rates of woody plants, herbs, and bryophytes were on average 2.71, 0.10, and 0.13 Mg ha−1; 0.24, 0.13, and 0.08 Mg ha−1 year−1 and 9.7%, 127.3%, and 67.6% year−1, respectively. Total understorey biomass and production increased continuously with stand age and reached the maximum in 146-year-old stands. Herbaceous biomass and production, peaked in 16-year-old stands, and remained stable thereafter with a slight increase in 146-year-old stands; in contrast, woody plant and bryophyte biomass and production peaked in 98- and 146-year-old stands. Herbaceous and woody vegetation turnover rates were higher in young stands, and those of bryophytes were higher in older stands. Total, woody and herbaceous biomass, production and turnover rates were higher under deciduous broadleaf overstorey, while values for bryophytes were higher in conifer stands, with mixedwood being intermediate. Overstorey mixture had no overall effect on total, woody, herbaceous biomass and production other than the expected from the averages of their broadleaf and conifer stands, but reduced bryophyte biomass and woody turnover. Our findings suggest that understorey biomass, production and turnover rates in the boreal forest are mainly driven by time since colonization and the resource availability and heterogeneity mediated via overstorey composition. Our results highlight the roles of stand development and overstorey composition in understorey biomass, production and turnover and their importance for modeling total forest ecosystem contribution to the global forest carbon.

Decreased occurrence of carbon cycle functions in microbial communities along with long-term secondary succession

The succession of microbial community structure and function is a central ecological topic; however, the mechanisms that underlie community assembly and promote temporal succession remain unclear. We studied microbial community-associated functional dynamics in a well-established secondary successional chronosequence that spans approximately 160 years of ecosystem development on the Loess Plateau of China, by sequencing both 16S and ITS rRNA genes and soil metagenomes, resulting in a total of 132.5 Gb of data. Notably, both bacterial and fungal communities shifted with succession, but the microbial community changed little from the pioneer forest stage (approximately 110 years) to the latter successional forest stage. Fungi showed higher variability with succession than bacteria, and the shift of both the bacterial and fungal communities was related more to the soil characteristics than to the litter characteristics. Shifts in soil microbial functions were associated with microbial phylogenetic changes, but microbial gene function also showed changes in the absence of phylogenetic changes at the late successional stages. The reduction in microbial C cycle genes was related to a decrease in litter decomposition ability, thus resulting in a steady state of nutrient cycle in the ecosystem. In addition, high microbial respiration in nutrient-rich soil does not necessarily indicate high microbial decomposition functions; the latter also depend on the abundance of related genes, on enzyme activity and on the physicochemical properties of the litter. Our study provides a metagenome profile of a successional chronosequence and provides insight into the mechanisms underlying the soil microbe-driven functional changes in nutrient cycles during succession.

Shade trees composition and diversity in cacao agroforestry systems of southern Pará, Brazilian Amazon

Agroforestry systems (AFS) are important agricultural land use in synergy with socio-environmental aspects, especially with cacao (Theobroma cacao L.) crop, a commodity mainly produced by smallholders in the humid tropics. In southern Para, Brazilian Amazon, farmers manage native shade trees growing with cacao, but species selection may be not appropriate to AFS maintenance over time. The objective of this study was to understand the shade trees transition between successional management phases of cacao-AFS, considering its initial shade (IS) and secondary shade (SS). It was sampled 10 plots in each situation (20,000 m2 in total) identifying individuals with CBH ≥ 15 cm. As expected, floristic composition was different and SS had greater species richness and diversity than IS, where only 17% of species were the shared among them. Musa sp. and Carica papaya L. were found only in IS and were dominant species, representing almost a half of the individuals. Although there was increase of late succession species from IS to SS, this still keeps high abundance of early succession species, such as Cecropia sp. The result shows an unexploited potential products and gap of services provision, such as N-fixing. The conclusion highlights the necessity of long-term succession planning and management practices to guarantee cacao crop maintenance and improve diversification with other income sources, such as fruits and wood. The role of biodiversity conservation, provided by shade trees, should be the target of political strategies to encourage its maintenance, such as payment for ecosystems services or other economic incentives.

Interaction between Ailanthus altissima and Native Robinia pseudoacacia in Early Succession: Implications for Forest Management

The goal of this study was to discover the nature and intensity of the interaction between an exotic invader Ailanthus altissima (Mill.) Swingle and its coexisting native Robinia pseudoacacia L. and consider management implications. The study occurred in the Mid-Appalachian region of the eastern United States. Ailanthus altissima can have a strong negative influence on community diversity and succession due to its allelopathic nature while R. pseudoacacia can have a positive effect on community diversity and succession because of its ability to fix nitrogen. How these trees interact and the influence of the interaction on succession will have important implications for forests in many regions of the world. An additive-replacement series common garden experiment was established to identify the type and extent of interactions between these trees over a three-year period. Both A. altissima and R. pseudoacacia grown in monoculture were inhibited by intraspecific competition. In the first year, A. altissima grown with R. pseudoacacia tended to be larger than A. altissima in monoculture, suggesting that R. pseudoacacia may facilitate the growth of A. altissima at the seedling stage. After the second year, R. pseudoacacia growth decreased as the proportion of coexisting A. altissima increased, indicating inhibition of R. pseudoacacia by A. altissima even though the R. pseudoacacia plants were much larger aboveground than the A. altissima plants. In early successional sites A. altissima should be removed, particularly in the presence of R. pseudoacacia in order to promote long-term community succession.

The influence of fire interval on community structure in the Greater Yellowstone Ecosystem

With fires becoming more common in the intermountain West, understanding patterns of post-fire succession and the role of fire interval in shaping community responses has become critical. In 2016, the Berry Fire burned through 4 study grids which have been the focus of a long-term fire succession project started after the massive 1988 fire season. We investigated the effects of this fire with respect to the repeatability of post-fire succession patterns (i.e., does succession after the 2016 fire match patterns observed after the 1988 fire) and the role of burn interval in shaping community structure of small mammals, invertebrates, and plants. Preliminary results indicate that sites with short burn intervals had greater diversity and abundance across all three taxonomic groups, although these differences were not always significant. Whereas the dominant taxa (deer mice and ants) were the same 1 year after the 1988 and 2016 fires, we documented greater abundance of invertebrates and mammals and greater diversity of mammals after the 2016 fires. Taken together, these results highlight the importance of understanding fire regime (i.e., fire timing and intensity) in shaping these ecological communities into the future.
 
 Featured photo by filarwilliams on Flickr. https://flic.kr/p/KYMiJF

Влияние доступности территории и характеристик рельефа на расположение и размер гарей в темнохвойных лесах Печоро-Илычского заповедника

Fires – one of the most important factors in catastrophic transformation of natural forest cover. Destroying whole ecosystems, fires, at the same time, start long-term succession processes. The study of the causes of fires makes a significant contribution to understanding structural and functional organization and dynamics of modern forests. The article examines the impact of availability of the territory and landscape-topographical features on the distribution and size of burnt areas formed in the last 150 years, in the foothill area of Pechora-Ilych Nature Reserve. The peculiarity of the territory is a complete lack of roads; the only roads are the rivers: Pechora, Ilych and their major tributaries. Analysis of the location of burned areas showed that significant influence on the occurrence of fire is provided by proximity of navigable rivers, since the greatest number of fires during studied period occurred at a distance of up to 3 km from navigable rivers, along which local population actively explores the territory (building of hunting shacks, gathering pine nuts, selective logging, etc.). Size of the fire is connected with proximity to navigable rivers: the largest fires is confined to the river banks and are likely to be only of anthropogenic origin. Small fires occurred at different distances from navigable rivers, could be both natural and anthropogenic ones. Settlements that existed before the formation of reserve had substantial and statistically significant effect on the occurrence of fire. Statistically significant associations of burnt areas with absolute altitude (the vast majority of burned areas are located on surfaces below 400 m) indirectly confirms that fires occurred at intensively developed parts of forests along the rivers.

Genesis and geomorphic evolution of the Velké pinky stopes in the Zlatohorská Highlands, Eastern Sudetes

Montanogenic landforms are commonly viewed as hazards by society, but they are also holders of specific and uncommon morphology with unique dynamics that act as remarks on landscape history. The Velké pinky stopes in the Zlatohorská Highlands, Eastern Sudetes, are naturally revitalized post-mining landforms with long-term geoecological succession. Their genetic origin is diverse but recent processes, such as deep-seated slope deformations, rockfalls and ground subsidence, have resulted in a distinct morphology dominated by rock walls and the accumulation of blocks and debris wedges. This morphology predisposes the stopes to become the core area of the most recent dynamic geomorphic activity within the wider, relatively homogenous area. By dendrogeomorphic techniques, we identified more than 20 rockfall events within three of the stopes with increased activity since the 1980s. Only the 1991 and 2006 events were identical for all three stopes. We obtained the years of exposures from 10 roots, revealing the ground subsidence and opening of tension cracks or even the lateral retreat of the flanks of minor depressions. The Schmidt hammer test revealed the most recent and fresh gravitational activity in one of the stopes. A relationship between the superficial morphology and underground structure was proven by electrical resistivity profiling. Compared to the previous studies engaged with the abandoned mines, we presented how complex their evolution can be. Our study brings new information about the historical development of anthropogenic relief forms. Moreover, our results suggest that standard research approaches can be successfully applied for development analysis of these specific forms.

Long-term succession of Collembola communities in relation to climate change and vegetation

The successional dynamics of Collembola communities on a rubble and debris dump in Bremen, Germany were studied between 1980 and 2000 in two experimental plots. One plot (SUC) was left undisturbed for natural secondary succession, the other plot (REC) was recultivated with rotary-tilling and sowing of grass. The mean Collembola abundance varied between 4 and 119 thousand ind. m−2 on SUC and between 6 and 92 thousand ind. m−2 on REC. In the two plots individual numbers peaked in the first decade, thereafter they decreased. Number of species varied from 9 to 17 on SUC with a very slight increase over time and from 8 to 17 on REC with only a temporary depression in 1998. The alpha diversity calculated as Shannon-Weaver and Simpson-indices increased in both plots. The lowest species turnover rate occurred between 1990 and 1994 on SUC and between 1981 and 1982 on REC, indicating the lowest diversity in species composition. During the whole succession, species composition changed more or less irregular in the two plots. With 74%, the dominance similarity (Renkonen index) of the Collembola communities between SUC and REC was greatest two years after the beginning of succession. As a result from CCA analysis, confirmed by linear regression analyses, Collembola total abundance and species composition (dominances) were significantly affected by annual (Oct-Sep) mean air and soil temperature (−10cm), increasing over the study period of twenty years. Ordination scores of Collembola species abundance on the CCA axes revealed four climatic groups of Collembola: cold-dry, cold-wet, warm-wet and warm-dry. A correlation of Collembola species abundance with vegetation cover was not found. Nevertheless, stronger effects of climatic variables were found on REC, which had a shorter vegetation than SUC, creating a different microclimate. Successional dynamics of Collembola communities in the studied plots are correlated with temporal shifts of species between early and late years of succession, but are also significantly influenced by soil temperature.

Eskişehir ilinde kırsal ve şehirsel habitatlardaki domuz leşlerine çekilen Coleoptera türlerinin süksesyon farklılıkları

Bu calismanin amaci, Eskisehir ilinde domuz lesi uzerindeki Coleoptera suksesyonunun belirlenmesi ve kirsal ve sehirsel habitatlardaki suksesyonal duzen farkliliklarinin degerlendirilmesidir. Eskisehir ilinde bulunan bir kirsal bir de sehirsel alana toplam 24 domuz lesi, Haziran 2012 ve Mayis 2013 arasinda bir yillik sure boyunca yerlestirilmistir. Coleoptera takimi icerisinde Staphylinidae, Histeridae, Dermestidae, Silphidae ve Cleridae familyalarina ait toplam 80 tur, ya direkt olarak les uzerinden ya da toprakta bulunan turler icin elemeyle toplanmistir. Tum turlerin aylari ve farkli curume evrelerinde bulunduklari sureler belirlenmistir. Buna ek olarak Turkiye’nin les Coleoptera faunasina yeni kayitlar eklenmistir. Yapilan calismada, cok yakin alanlarda bile hem tur sayisinin hem de aktivite periyodlarinin belirgin sekilde farklilik gosterdigi aciga cikmaktadir. Bu sekildeki uzun-sureli suksesyon calismalarinin Turkiye’nin tum illerinde yapilmasi gerekliligi ortaya konulmustur. Elde edilen veriler Turkiye’de adli olaylarda minimum olum zaman araligi tahmininde potansiyel olarak kullanilabilecektir.

Vegetation Succession and Environmental Conditions following Catastrophic Lake Drainage in Old Crow Flats, Yukon

Increases in the frequency and magnitude of disturbances associated with the thawing of ice-rich permafrost highlight the need to understand long-term vegetation succession in permafrost environments. This study uses field sampling and remote sensing to explore vegetation development and soil conditions following catastrophic lake drainage in Old Crow Flats (OCF). The data presented show that vegetation on drained lake basins in OCF is characterized by two distinct assemblages: tall willow stands and sedge swards. Field sampling indicates that these alternative successional trajectories result from variation in soil moisture following drainage. Increased willow mortality on older drained basins suggests that intraspecific competition drives self-thinning in shrub thickets. This finding, combined with data from paleoecological studies and contemporary vegetation in OCF, suggests that willow stands on drained lake basins are seral communities. These results also indicate that the increase in number of catastrophic drainages that occurred between 1972 and 2010 will alter regional vegetation in ways that affect wildlife habitat, permafrost conditions, and local hydrology.

A foundation of ecology rediscovered: 100years of succession on the William S. Cooper plots in Glacier Bay, Alaska.

Understanding plant community succession is one of the original pursuits of ecology, forming some of the earliest theoretical frameworks in the field. Much of this was built on the long-term research of William S. Cooper, who established a permanent plot network in Glacier Bay, Alaska, in 1916. This study now represents the longest-running primary succession plot network in the world. Permanent plots are useful for their ability to follow mechanistic change through time without assumptions inherent in space-for-time (chronosequence) designs. After 100-yr, these plots show surprising variety in species composition, soil characteristics (carbon, nitrogen, depth), and percent cover, attributable to variation in initial vegetation establishment first noted by Cooper in the 1916-1923 time period, partially driven by dispersal limitations. There has been almost a complete community composition replacement over the century and general species richness increase, but the effective number of species has declined significantly due to dominance of Salix species which established 100-yr prior (the only remaining species from the original cohort). Where Salix dominates, there is no establishment of "later" successional species like Picea. Plots nearer the entrance to Glacier Bay, and thus closer to potential seed sources after the most recent glaciation, have had consistently higher species richness for 100yr. Age of plots is the best predictor of soil N content and C:N ratio, though plots still dominated by Salix had lower overall N; soil accumulation was more associated with dominant species. This highlights the importance of contingency and dispersal in community development. The 100-yr record of these plots, including species composition, spatial relationships, cover, and observed interactions between species provides a powerful view of long-term primary succession.

Многолетняя сукцессия фитопланктона среднего течения реки Иртыш (Омск, Россия)

Многолетняя сукцессия фитопланктона среднего течения реки Иртыш (Омск, Россия)

Top-down vs. bottom-up control on vegetation composition in a tidal marsh depends on scale

The relative impact of top-down control by herbivores and bottom-up control by environmental conditions on vegetation is a subject of debate in ecology. In this study, we hypothesize that top-down control by goose foraging and bottom-up control by sediment accretion on vegetation composition within an ecosystem can co-occur but operate at different spatial and temporal scales. We used a highly dynamic marsh system with a large population of the Greylag goose (Anser anser) to investigate the potential importance of spatial and temporal scales on these processes. At the local scale, Greylag geese grub for below-ground storage organs of the vegetation, thereby creating bare patches of a few square metres within the marsh vegetation. In our study, such activities by Greylag geese allowed them to exert top-down control by setting back vegetation succession. However, we found that the patches reverted back to the initial vegetation type within 12 years. At large spatial (i.e. several square kilometres) and temporal scales (i.e. decades), high rates of sediment accretion surpassing the rate of local sea-level rise were found to drive long-term vegetation succession and increased cover of several climax vegetation types. In summary, we conclude that the vegetation composition within this tidal marsh was primarily controlled by the bottom-up factor of sediment accretion, which operates at large spatial as well as temporal scales. Top-down control exerted by herbivores was found to be a secondary process and operated at much smaller spatial and temporal scales.

Long-Term Succession of the Phytoplankton of the Irtysh River Middle Flow (Omsk, Russia)

Long-Term Succession of the Phytoplankton of the Irtysh River Middle Flow (Omsk, Russia)

Modeling long-term changes in tundra carbon balance following wildfire, climate change, and potential nutrient addition.

To investigate the underlying mechanisms that control long-term recovery of tundra carbon (C) and nutrients after fire, we employed the Multiple Element Limitation (MEL) model to simulate 200-yr post-fire changes in the biogeochemistry of three sites along a burn severity gradient in response to increases in air temperature, CO2 concentration, nitrogen (N) deposition, and phosphorus (P) weathering rates. The simulations were conducted for severely burned, moderately burned, and unburned arctic tundra. Our simulations indicated that recovery of C balance after fire was mainly determined by the internal redistribution of nutrients among ecosystem components (controlled by air temperature), rather than the supply of nutrients from external sources (e.g., nitrogen deposition and fixation, phosphorus weathering). Increases in air temperature and atmospheric CO2 concentration resulted in (1) a net transfer of nutrient from soil organic matter to vegetation and (2) higher C : nutrient ratios in vegetation and soil organic matter. These changes led to gains in vegetation biomass C but net losses in soil organic C stocks. Under a warming climate, nutrients lost in wildfire were difficult to recover because the warming-induced acceleration in nutrient cycles caused further net nutrient loss from the system through leaching. In both burned and unburned tundra, the warming-caused acceleration in nutrient cycles and increases in ecosystem C stocks were eventually constrained by increases in soil C : nutrient ratios, which increased microbial retention of plant-available nutrients in the soil. Accelerated nutrient turnover, loss of C, and increasing soil temperatures will likely result in vegetation changes, which further regulate the long-term biogeochemical succession. Our analysis should help in the assessment of tundra C budgets and of the recovery of biogeochemical function following fire, which is in turn necessary for the maintenance of wildlife habitat and tundra vegetation.

Response of phytoplankton to protective-restoration treatments enhancing water quality in a shallow urban lake.

Lake Jeziorak Mały is a shallow urban lake where storm water pretreatment separators and a fountain-based water aeration system were installed as protective-restoration measures to enhance water quality. We investigated the effect of these procedures on phytoplankton dynamics and physicochemical properties in the littoral and pelagial zones in 1996–2003, 2005, and 2013. A decrease in cyanobacteria proportion, abundance, and biomass has been noticed, and other phytoplankton groups increased after these procedures. Significantly elevated species diversity was recorded in the littoral zone with the exchange of cyanobacteria and diatom dominant species typically induced by alteration from hypertrophic to eutrophic status. For example, the polytrophic Limnothrix redekei was replaced by eutrophic Planktolyngbya brevicellularis. This stemmed from greater oxygenation, water visibility and diminished pH, conductivity, and orthophosphates. Our results showed that introducing these restoration measures influence on the long-term succession of phytoplankton and induced the change from a polytrophic to eutrophic state, and that such measures are vitally important in future considerations of shallow urban lake management.

The change of soil water storage in three land use types after 10 years on the Loess Plateau

Soil water content (SWC) is a critical variable in studies of hydrological processes and the soil–plant–atmosphere continuum, especially in arid and semi-arid regions such as the Loess Plateau. Knowledge of the effects of vegetation types on soil water dynamics would aid in understanding the mechanisms responsible for water shortages and addressing the problem of poor long-term vegetation recovery. To evaluate the response of soil water storage (SWS) to different land use types during long-term natural vegetation succession, we examined soil moisture under different land use types (grassland, shrubland and forestland) at different times (2005 and 2014) in the Ziwuling forest region, located in the central part of the Loess Plateau, China. Our results showed that vegetation type had a significant effect on SWS in the 0–500cm soil layer. The SWS in grassland and shrubland was significantly higher in August 2014 than in August 2005, but the pattern was reversed for forestland. In the same year, the SWS was highest in grassland, intermediate in shrubland, and lowest in forestland. SWS and SWC showed a highly significant and positive relationship in the 0–60cm soil layers (P<0.01), and both soil bulk density (BD) and soil total nitrogen (TN) showed a significant relationship with SWS. In addition, the SWC and soil total porosity (TP) showed a significant positive relationship in the 0–20cm soil layer (P<0.05). These results are expected to provide insights into land water management and inform efforts to estimate the productivity and sustainability of semiarid ecosystems.

Succession of arthropods on xerothermophilous habitats formed by sand quarrying: Epigeic beetles (Coleoptera) and orthopteroids (Orthoptera, Dermaptera and Blattodea)

European Commision suggested the remedying biodiversity damage should be achieved through restoration of the environment as a whole to its baseline condition. Here we provide the evidence on non-market goods gained when spontaneous succession is allowed at sites subject to previous sand or gravel-sand quarrying, arguing for estimation of the value of natural resources formed during the quarrying process and for their reflection in the restoration plans. We analyzed the assemblages of beetles and orthopteroids formed at xerothermophilous microhabitats subject to spontaneous succession for 2–5, 6–15 and >15years, or formed at reclaimed sites within sandpits and gravel-sandpits in the Czech Republic, Central Europe. We collected a total of 7259 individuals from 401 species of beetles, and 2785 individuals from 43 species of orthopteroids. We found 62 red-listed species, among them eight CR species, including abundant Psammodius asper and Zorochros meridionalis; Labidura riparia (EN) was eudominant orthopteroid of early successional stages. Xerothermes subject to spontaneous succession hosted 95% of red-listed species found, whereas reclaimed sites hosted mostly the generalists and only 24% of red-listed species found. We identified 13 species specialized for drift sand (all beetles); most were present at xerothermes subject to intermediate or long-term succession. We identified 53 beetle and five orthopteroid species specialized for gravel-sand river terraces and xerothermophilous steppes. Striking 97% of red-listed beetles and 80% of red-listed orthopteroids found in the sandpits were absent in other comparable postindustrial habitats with fine-grained substrates. Sandpit xerothermes thus serve as important secondary strongholds for psammophilous beetles and orthopteroids.

Patterns of taxonomic, phylogenetic diversity during a long-term succession of forest on the Loess Plateau, China: insights into assembly process.

Quantifying the drivers underlying the distribution of biodiversity during succession is a critical issue in ecology and conservation, and also can provide insights into the mechanisms of community assembly. Ninety plots were established in the Loess Plateau region of northern Shaanxi in China. The taxonomic and phylogenetic (alpha and beta) diversity were quantified within six succession stages. Null models were used to test whether phylogenetic distance observed differed from random expectations. Taxonomic beta diversity did not show a regular pattern, while phylogenetic beta diversity decreased throughout succession. The shrub stage occurred as a transition from phylogenetic overdispersion to clustering either for NRI (Net Relatedness Index) or betaNRI. The betaNTI (Nearest Taxon Index) values for early stages were on average phylogenetically random, but for the betaNRI analyses, these stages were phylogenetically overdispersed. Assembly of woody plants differed from that of herbaceous plants during late community succession. We suggest that deterministic and stochastic processes respectively play a role in different aspects of community phylogenetic structure for early succession stage, and that community composition of late succession stage is governed by a deterministic process. In conclusion, the long-lasting evolutionary imprints on the present-day composition of communities arrayed along the succession gradient.