The changes in soil organic carbon stock and quality across a subalpine forest successional series

Soil organic carbon(SOC)affects the function of terrestrial ecosystem and plays a vital role in global carbon cycle.Yet,large uncertainty still existed regarding the changes in SOC stock and quality with forest succession.Here,the stock and quality of SOC at 1-m soil profile were investigated across a subalpine forest series,including shrub,deciduous broad-leaved forest,broadleaf-conifer mixed forest,middle-age coniferous forest and mature coniferous forest,which located at southeast of Tibetan Plateau.The results showed that SOC stock ranged from 9.8 to29.9 kg·m^(-2),and exhibited a hump-shaped response pattern across the forest successional series.The highest and lowest SOC stock was observed in the mixed forest and shrub forest,respectively.The SOC stock had no significant relationships with soil temperature and litter stock,but was positively correlated with wood debris stock.Meanwhile,the average percentages of polysaccharides,lignins,aromatics and aliphatics based on FTIR spectroscopy were 79.89%,0.94%,18.87%and 0.29%,respectively.Furthermore,the percentage of polysaccharides exhibited an increasing pattern across the forest successional series except for the sudden decreasing in the mixed forest,while the proportions of lignins,aromatics and aliphatics exhibited a decreasing pattern across the forest successional series except for the sudden increasing in the mixed forest.Consequently,the humification indices(HIs)were highest in the mixed forest compared to the other four successional stages,which means that the SOC quality in mixed forest was worse than other successional stages.In addition,the SOC stock,recalcitrant fractions and HIs decreased with increasing soil depth,while the polysaccharides exhibited an increasing pattern.These findings demonstrate that the mixed forest had higher SOC stock and worse SOC quality than other successional stages.The high proportion of SOC stock(66%at depth of 20-100 cm)and better SOC quality(lower HIs)indicate that deep soil have tremendous potential to store SOC and needs more attention under global chan ge.

Changes in plant debris and carbon stocks across a subalpine forest successional series

Background:As a structurally and functionally important component in forest ecosystems,plant debris plays a crucial role in the global carbon cycle.Although it is well known that plant debris stocks vary greatly with tree species composition,forest type,forest origin,and stand age,simultaneous investigation on the changes in woody and non-woody debris biomass and their carbon stock with forest succession has not been reported.Therefore,woody and non-woody debris and carbon stocks were investigated across a subalpine forest successional gradient in Wanglang National Nature Reserve on the eastern Qinghai-Tibet Plateau.Results:Plant debris ranged from 25.19 to 82.89 Mg∙ha−1 and showed a global increasing tendency across the subalpine forest successional series except for decreasing at the S4 successional stage.Accordingly,the ratios of woody to non-woody debris stocks ranged from 26.58 to 208.89,and the highest and lowest ratios of woody to non-woody debris stocks were respectively observed in mid-successional coniferous forest and shrub forest,implying that woody debris dominates the plant debris.In particular,the ratios of coarse to fine woody debris stocks varied greatly with the successional stage,and the highest and lowest ratios were found in later and earlier successional subalpine forests,respectively.Furthermore,the woody debris stock varied greatly with diameter size,and larger diameter woody debris dominated the plant debris.Correspondingly,the carbon stock of plant debris ranged from 10.30 to 38.87 Mg∙ha−1 across the successional series,and the highest and lowest values were observed in the mid-coniferous stage and shrub forest stage,respectively.Most importantly,the carbon stored in coarse woody debris in later successional forests was four times higher than in earlier successional forests.Conclusions:The stock and role of woody debris,particularly coarse woody debris,varied greatly with the forest successional stage and dominated the carbon cycle in the subalpine forest ecosystem.Thus,preserving coarse woody debris is a critical strategy for sustainable forest management.

Variations of bacterial and fungal communities along a primary successional chronosequence in the Hailuogou glacier retreat area(Gongga Mountain, SW China)

New terrestrial habitats have emerged and a primary succession has developed in the retreat area （29°34＇N, 102°oo＇E, 2951-2886 m） after the retreat of the Hailuogou glacier. To investigate soil microbial changes along the primary successional chronosequence, mixed soil samples were collected at six sites at different ages （2 young sites, 2 mid-aged sites, and 2 old sites）. The RNA was extracted and amplified. Bacterial 16S rRNA and fungal 18S rRNA were analyzed using high-throughput 454 pyrosequencing analysis. Overall, pyrosequeneing showed that Proteobacteria, Acidobacteria, Baeteroidetes and Actinobacteria were the main bacterial phyla, and the fungal communities were strongly dominated by the phyla Ascomyeota and Basidiomyeota in the retreat area. The Shannon diversity index （Hshannon） of bacteria was 6.5 - 7.9, and that of fungi was 2.2 - 4.1 in these sites. For the bacterial communities, diversity and evenness values were highest on the mid-age sites and were relatively low on the young trend was observed for the and old sites. A similar fungal communities. In contrast, soil properties showed significant linear distributional trends （increase or decrease） with the age of the site. Combining the linear change patterns of soil properties, the highest values of bacterial and fungal evenness and diversity in the mid-aged sites indicated that there was less environmental stress and more niches for microbial communities in the middle successional stage compare with other stages. In addition, our analysis showed that microbial communities were the main drivers that build a soil organic matter pool to expedite pedogenesis for ecosystem succession. This primary succession in the Hailuogou glacier retreat area is developing rapidly compared with that in other glacier retreats.

Peashrub Community May Accelerate the Successional Process in a Meadowland-Peashrub-Birch Sere

Net N mineralization and nitrification were determined using the closed_top PVC tube in situ incubation method in a subalpine meadow (Saussurea iodostegia Hance + Carex capillaris L.)→shagspine peashrub (Caragana jubata (Pall.) Poir) shrubland→ribbed birch (Betula costata Trautv.) successional sere. The ability of the three communities to supply available N was comparatively studied. The results showed that there were apparent seasonal changes in the inorganic N pools (including NH + 4_N and NO - 3_N) and net N mineralization and nitrification rates in the three sites. There were generally no significant differences in the inorganic N pools among sampling events. But the NH + 4_N concentration in both birch (P<0.01) and meadow (P<0.01) sites was significantly higher than that in peashrub site in June 1996, and the NO - 3_N concentration in peashrub site was significantly higher than that of meadow site (P<0.05) in August 1996. The annual net N mineralization and nitrification rates in peashrub site (16.01 kg·hm -2 ) were higher than in birch (12.05 kg·hm -2 ) and meadow sites (1.64 kg·hm -2 ). The annual net nitrification rate in peashrub site (11.37 kg·hm -2 ) was higher than in meadow site (10.90 kg·hm -2 ) and much lower than in birch site (14.36 kg·hm -2 ). We conclude that the ability of peashrub shrubland soil to supply available N for plant uptake and the ability to prevent available N from denitrification and leaching were higher than that of the other two sites. The leguminous peashrub might play a potential role in supplying more N, which in turn facilitate the invasion of birch saplings during the successional processes.

Integrating species and successional classes for wood production in a mixed forest restoration system in a neotropical region

As forests in neotropical regions,particularly in developing countries,are devastated,interventions to restore biodiversity and its ecological functions are needed.Rural producers have thus been encouraged to grow trees for wood production as an economic activity.The objective of this study was to quantify the increment of wood density of four forest species from different successional classes of a mixed system of restoration in a neotropical forest in Brazil.Tree discs were sampled at breast height and analyzed radially by X-ray densitometry to obtain apparent density and basic density.Three trees each of a species from the pioneer,early and late secondary successional stages and of the dominant species in the climax community.The radial profiles indicated increasing density from the pith to bark of trunks,except for some variations due to wood defects and growth.Average density was 0.576 g cm^(-3),0.655 g cm^(-3),0.706 g cm^(-3)and 0.775 g cm^(-3),respectively,for Peltophorum dubium,Schinus terebinthifolius,Cariniana estrellensis and Hymenaea courbaril.Radial profiles indicated higher amplitudes in the apparent densities for slow-growing species.X-ray densitometry generated parameters such as minimum,average and maximum densities,and radial density variations.These parameters are important for understanding the ecological functional role of successional classes of the Atlantic Forest from the Neotropical region.

Understorey species distinguish late successional and ancient forests after decades of minimum human intervention: A case study from Slovenia

The main species composition drivers in temperate deciduous forests are environmental conditions, a stand?s age and the site history, e.g., the succession stage and past land use, as well as disturbance regime and current management. We compared plant species diversity and composition in late successional and ancient forests, cooccurring on the same small river island applying species accumulation curves and nonmetric multidimensional scaling, respectively. Given the island?s geomorphological characteristics, we expected these to be very similar before human intervention in the past. The forests experienced differing disturbance regimes in the past, while over the last 30 years, human intervention has been the same and reduced to a minimum. The ancient forest in this study had two major characteristics defining it as old, mature forest: continuity of presence for more than 200years and specific composition. The late successional forest experienced major disturbance in the 20th century and was allowed natural regeneration by bordering on the ancient forest, representing a potential species pool,and by decades of minimum human intervention. Our results showed that, even though there was no difference in species richness, we could still detect differences between the forests, particularly in the abundance and species composition of the understorey, among which geophytes had the most indicative importance. To make our results useful on a broader scale, we composed from the literature a species list of plants indicative for ancient forest and tested its application. These results are important for distinguishing between old and mature secondary stands and particularly for identifying old forest stands, which should be conserved and, in the case of fragmented landscapes, included in a network connecting forest fragments.

Changes of Soil Water, Organic Matter, and Exchangeable Cations Along a Forest Successional Gradient in Southern China

Information on the distribution patterns of soil water content （SWC）, soil organic matter （SOM）, and soil exchangeable cations （SEC） is important for managing forest ecosystems in a sustainable manner. This study investigated how SWC, SOM, and SEC were influenced in forests along a successional gradient, including a regional climax （monsoon evergreen broad-leaved forest, or MEBF）, a transitional forest （coniferous and broad-leaved mixed forest, or MF）, and a pioneer forest （coniferous Masson pine （Pinus rnassoniana） forest, or MPF） of the Dinghushan Biosphere Reserve in the subtropical region of southern China. SWC, SOM, and SEC excluding Ca^2＋ were found to increase in the soil during forest succession, being highest in the top soil layer （0 to 15 cm depth） except for Na^＋. The differences between soil layers were largest in MF. This finding also suggested that the nutrients were enriched in the topsoil when they became increasingly scarce in the soil. There were no significant differences （P = 0.05） among SWC, SOM, and SEC. A linear, positive correlation was found between SWC and SOM. The correlation between SOM and cation exchange capacity （CEC） was statistically significant, which agreed with the theory that the most important factor determining SEC is SOM. The ratio of K^＋ to Na^＋ in the topsoil was about a half of that in the plants of each forest. MF had the lowest exchangeable Ca^2＋ concentration among the three forests and Ca^2＋：K^＋ in MPF was two times higher than that in MF. Understanding the changes of SWC, SOM, and CEC during forest succession would be of great help in protecting all three forests in southern China.

Regenerative Role of Soil Seed Banks of Different Successional Stages in A Saline-alkaline Grassland in Northeast China

Soil seed banks can act as a potential seed source for natural revegetation and restoration. However, in a saline-alkaline grassland, it remains unclear how the stages of vegetation succession affect the characteristics of soil seed banks and the potential of soil seed banks of different successional stages for vegetation restoration. In this study, seasonal changes of the soil seed bank, and seed production and dispersal dynamics along degradation successional gradients were investigated in a saline-alkaline grassland in Northeast China, where the dominant grass during the 1960 s, Leymus chinensis was replaced with the secondary successional order of Puccinellia chinampoensis, Chloris virgata, and Suaeda salsa, together with bare patches. It was found that the soil seed bank composition varied according to the changing vegetation and had the highest species richness(7–16) in the climax successional stage, but had a low S?rensen similarity(0.22–0.37) with the aboveground vegetation. There was a high seed density of the soil seed bank(21 062–62 166/m2 in August and December) and also high S?rensen similarity index values(0.47–0.60) in the secondary successional stages of P. chinampoensis, C. virgata, and S. salsa. In bare patches, there were many seeds in the soil seed bank and some seedlings also appeared in the aboveground vegetation, indicating the existence of a persistent soil seed bank. Seed density and species richness differed substantially among the different successional stages, which was related to the reproductive characteristics of the standing plants in vegetation communities. Due to the lack of propagules of perennial species, especially the climax species of L. chinensis, in the soil, the successful restoration of the degraded saline-alkaline grassland was not possible. The study proved that in a degraded saline-alkaline grassland dominated by biennial or annual species, the soil seed bank was important for the revegetation of the current dominant plants, but not for the restoration of the original target species. Therefore, it is necessary to induce seeds or other propagules of the target perennial species.

Changes in microbial community composition in the leaf litter of successional communities after volcanic eruptions of Mount Usu, northern Japan

Changes in the fungal and bacterial biomass and community structure in litter after the volcanic eruptions of Mount Usu, northern Japan were investigated using a chronosequence approach, which is widely used for analyzing vegetation succession. The vegetation changed from bare ground （10 years after the eruptions） with little plant cover and poor soil to monotonic grassland dominated by Polygonum sachalinense with undeveloped soil （33 years） and then to deciduous broad-leaved forest dominated by Populus maximowiczii with diverse species composition and well-developed soil （100 years）. At three chronosequential sites, we evaluated the compositions of phospholipid fatty acids （PLFAs）, carbon （C） and nitrogen （N） contents and the isotope ratios of C （δ13C） and N （δ15N） in the litter of two dominant species, Polygonum sachalinense and Populus maximowiezii. The C/N ratio, δ13C and δ15N in the litter of these two species were higher in the forest than that in the bare ground and grassland. The PLFAs gradually increased from the bare ground to the forest, showing that microbial biomass increased with the development of the soil and/or vegetation. The fungi-to-bacteria ratio of PLFA was constant at 5.3 ± 1.4 in all three sites, suggesting that fungi were predominant. A canonical correspondence analysis suggested that the PLFA comoosition was related to the successional ages and the developing soil properties （P 〈 0.05, ANOSIM）. The chrono- sequential analysis effectively detected the successional changes in both microbial and plant communities.

Assessing stand structure in successional stages of dark coniferous forests in western Sichuan, China

Subalpine dark coniferous forests in the western Sichuan Province of China play an important role in the hydrological processes in the upper reaches of the Yangtze River. Second-growth forests, with different stand successional stages, have developed as a result of logging over the past 50 years. Forest cover and stand structure changed greatly with concomitant degradation of forest ecosystem functions. To understand how the stand structures of the second-growth forests change during the stand succession process, we analyzed stand structure characteristics and an old-growth state index of the bamboo and moss-forest types. We found that stand structure at the young successional stage featured one-third of the structure characteristics of the old-growth dark coniferous forests,while the structure of the medium-aged stage had reached half the structure of the old-growth state. The two forest types were similar in the rate of development at the young successional stage but differed at the medium-aged stage;the moss-forest type had more advanced development than the bamboo-forest type at the medium-aged successional stage.

The carbon fluxes in different successional stages:modelling the dynamics of tropical montane forests in South Ecuador

Background： Tropical forests play an important role in the global carbon（C） cycle.However, tropical montane forests have been studied less than tropical lowland forests, and their role in carbon storage is not well understood.Montane forests are highly endangered due to logging, land-use and climate change.Our objective was to analyse how the carbon balance changes during forest succession.Methods： In this study, we used a method to estimate local carbon balances that combined forest inventory data with process-based forest models.We utilised such a forest model to study the carbon balance of a tropical montane forest in South Ecuador, comparing two topographical slope positions（ravines and lower slopes vs upper slopes and ridges）.Results： The simulation results showed that the forest acts as a carbon sink with a maximum net ecosystem exchange（NEE） of 9.3 Mg C？（ha？yr）-1during its early successional stage（0–100 years）.In the late successional stage, the simulated NEE fluctuated around zero and had a variation of 0.77 Mg C？（ha？yr）–1.The simulated variability of the NEE was within the range of the field data.We discovered several forest attributes（e.g., basal area or the relative amount of pioneer trees） that can serve as predictors for NEE for young forest stands（0–100 years） but not for those in the late successional stage（500–1,000 years）.In case of young forest stands these correlations are high, especially between stand basal area and NEE.Conclusion： In this study, we used an Ecuadorian study site as an example of how to successfully link a forest model with forest inventory data, for estimating stem-diameter distributions, biomass and aboveground net primary productivity.To conclude, this study shows that process-based forest models can be used to investigate the carbon balance of tropical montane forests.With this model it is possible to find hidden relationships between forest attributes and forest carbon fluxes.These relationships promote a better understanding of the role of tropical montane forests in the context of global carbon cycle, which in future will become more relevant to a society under global change.

Belowground carbon balance and carbon accumulation rate in the successional series of monsoon evergreen broad-leaved forest

The balance, accumulation rate and temporal dynamics of belowground carbon in the successional series of monsoon evergreen broadleaved forest are obtained in this paper, based on long-term observations to the soil organic matter, input and standing biomass of litter and coarse woody debris, and dissolved organic carbon carried in the hydrological process of subtropical climax forest ecosystem—monsoon evergreen broad-leaved forest, and its two successional forests of natural restoration—coniferous and broad-leaved mixed forest and Pinus massoniana forest, as well as data of root biomass obtained once every five years and respiration measurement of soil, litter and coarse woody debris respiration for 1 year. The major results include: the belowground carbon pools of monsoon evergreen broad-leaved forest, coniferous and broad-leaved mixed forest, and Pinus massoniana forest are 23191 ± 2538 g·m?2, 16889 ± 1936 g·m?2 and 12680 ± 1854 g·m?2, respec- tively, in 2002. Mean annual carbon accumulation rates of the three forest types during the 24a from 1978 to 2002 are 383 ± 97 g·m?2·a?1, 193 ± 85 g·m?2·a?1 and 213 ± 86 g·m?2·a?1, respectively. The belowground carbon pools in the three forest types keep increasing during the observation period, suggesting that belowground carbon pools are carbon sinks to the atmosphere. There are seasonal variations, namely, they are strong carbon sources from April to June, weak carbon sources from July to September; while they are strong carbon sinks from October to November, weak carbon sinks from December to March.

Dependence of Soil Respiration on Soil Temperature and Soil Moisture in Successional Forests in Southern China

The spatial and temporal variations in soil respiration and its relationship with biophysical factors In forests near the Tropic of Cancer remain highly uncertain. To contribute towards an Improvement of actual estimates, soil respiration rates, soil temperature, and soil moisture were measured In three successional subtropical forests at the Dlnghuahan Nature Reserve （DNR） In southern China from March 2003 to February 2005. The overall objective of the present study was to analyze the temporal variations of soil respiration and Its biophysical dependence in these forests. The relationships between biophysical factors and soil respiration rates were compared In successional forests to test the hypothesis that these forests responded similarly to biophysical factors. The seasonality of soil respiration coincided with the seasonal climate pattern, with high respiration rates in the hot humid season （April-September） and with low rates In the cool dry season （October-March）. Soil respiration measured at these forests showed a clear Increasing trend with the progressive succession. Annual mean （± SD） soil respiration rate In the DNR forests was （9.0 ± 4.6） Mg CO2-C/hm^2 per year, ranging from （6.1 ± 3.2） Mg CO2-C/hm^2 per year in early successional forests to （10.7 ± 4.9） Mg CO2-C/hm^2 per year in advanced successional forests. Soil respiration was correlated with both soil temperature and moisture. The T/M model, where the two biophysical variables are driving factors, accounted for 74%-82% of soil respiration variation In DNR forests. Temperature sensitivity decreased along progressive succession stages, suggesting that advanced-successional forests have a good ability to adjust to temperature. In contrast, moisture Increased with progressive succession processes. This increase is caused, in part, by abundant respirators In advanced-successional forest, where more soil moisture is needed to maintain their activities.

High-nitrogen and low-irradiance can restrict energy utilization in photosynthesis of successional tree species in low subtropical forest

Responses of photosynthesis and the partition of energy utilization to high-nitrogen importation and high-light intensity in leaves of three dominant tree species of subtropical forest,including sun plant or early-successional species Schima superba,mesophyte or intermediate-successional species Canstanopsis hystrix,and shading-tolerant plant or late-successional species Cryptocarya concinna were studied by using the CO2 exchange system and chlorophyll fluorescence method.Our results showed that,regardless of plant species,net photosynthetic rate(Pn)was higher in high-nitrogen supply and high irradiance(HNHL)plants than in low-nitrogen supply and high irradiance(LNHL)plants,implying that low-nitrogen importation would limit Pn of plants grown under high irradiance.However, high-nitrogen supply and low irradiance(HNLL)plants had a lower Pn.Insignificant change of quantum yield(Fv′/Fm′)in opened PS II was found in leaves of HNHL,LNHL or HNLL plants of S.superba and C. hystrix,while a higher Fv′/Fm′occurred in HNHL plants of C.concinna in comparison with LNHL or HNLL plants.The HNHL plants of C.concinna also had a higher photochemical quantum yield(△F/Fm′) than LNHL or HNLL plants,however no similar responses were found in plants of S.superba and C. hystrix(P<0.05).In the irradiance range of 0―2000μmol photon·m -2·s -1,the fraction of energy consumed by photochemistry(φ PSII )was 18.2%in LNHL plants of S.superba which was higher than that in HNHL plants(P>0.05)and it was significantly higher than in HNLL plants(P<0.05).C.hystrix also had a similar response inφ PSII to nitrogen supply and irradiance.Regardless of species HNLL plants had a significantφ PSII and higher heat dissipation in light,and this effect was more severe in C.concinna than in S.superba or C.hystrix.The results may mean that high-nitrogen importation by nitrogen deposit and low irradiance caused by changing climate or air pollution would more severely restrict photosynthetic processes in the late-successional species C.concinna than in the early-successional species S.superba and intermediate-successional species C.hystrix.The continuous high-nitrogen precipitation in the future and the over cast mist or pollution smoke could induce late-successional species to degrade,however,early-successional species would be more adapted to competition for more resources to keep their dominance in ecosystems.In this sense,the zonal vegetation may accelerate degradation caused by high nitrogen precipitation and low irradiance,while the early-successional and mesophytic vegetations can remain longer.Thus,nitrogen precipitation may play an important role in plant community succession.

Positive effects of tree species diversity on litterfall quantity and quality along a secondary successional chronosequence in a subtropical forest

Aims Litterfall,as an important link between aboveground and belowground processes,plays a key role in forest ecosystems.Here,we test for effects of tree species richness on litter production and litter quality in subtropical forest.The study further encompasses a factorial gradient of secondary succession that resulted from human exploitation.Given that a large percentage of subtropical forests are in secondary successional stages,understanding the role of biodiversity on forest re-growth after disturbance appears critical.Methods From January 2009 to December 2014,we monitored forest litterfall in 27 Comparative Study Plots that spanned a gradient of tree species richness(3-20 species)and secondary successional ages(~20 to 120 years)in Gutianshan Natural Nature Reserve,Zhejiang Province,China.The experiment is part of the biodiversity-ecosystem functioning research platform‘BEF-China’.Tree litterfall was collected in monthly intervals using litter traps.Samples were separated into leaf and non-leaf components.Leaf litter was further sorted into dominant and other species.Community level monthly leaf litter C and N contents were analysed through a full year.General linear mixed-effects models were applied to test for effects of tree species richness and successional age on litter quantity and leaf litter C/N.Important Findings Litterfall increased with species richness among and within successional age and this effect was consistent across years.Successionally older stands had higher litterfall and this effect was related to increased tree species richness.However,species richness did not change the intra-and inter-annual temporal stability of litterfall.Increasing tree species richness increased leaf litter quality(decreased C/N),while successional age had no effect.Our study indicates that more diverse forest stands produce more leaf litter and that this litter has higher N concentrations,which could promote forest growth through accelerated nutrient re-cycling.

Ramet Population Structure of Fargesia nitida （Mitford） Keng f. et Yi in Different Successional Stands of the Subalpine Coniferous Forest in Wolong Nature Reserve

Forest structure and succession in Wolong Nature Reserve is influenced by the understory dwarf bamboo population. However, less is known about how the forest succession affects the dwarf bamboo population. To examine the bamboo ramet population growth of Fargesla nitida （Mltford） Keng f. et Yi and to determine how ramet population structure varies along the succession of coniferous forest, we sampled ramet populations of F. nitida from the following three successional stages： （i） a deciduous broad-leaved （BL） stand; （ii） a mixed broad-leaved coniferous （MI） stand; and （ill） a coniferous （CF） stand. We investigated the population structure, biomass allocation, and morphological characteristics of the bamboo ramet among the three stand types. Clonal ramets, constituting the bamboo population, tended to become short and small with succession. The ramet changed towards having a greater mass investment in leaves, branches and underground roots and rhizomes rather than in the culm. With respect to leaf traits, individual leaf mass and area in the BL stand were markedly bigger than those In both the MI and CF stands, except for no significant difference in specific leaf area. The age distribution showed that the bamboo population approached an older age with succession. The results demonstrate that the ramet population structure of F. nitida is unstable and its growth performance is inhibited by succession.

Successional trends and processes on a glacial foreland in Southern Iceland studied by repeated species counts

Introduction:Primary succession on glacial forelands is increasingly relevant as rapid glacial retreat is exposing growing land areas to plant colonization.We investigated temporal trends,controls,and outcomes in floral succession on a subarctic glacial foreland.Specifically,we examined changes in community composition(mosses,low shrubs,forbs,trees,and graminoids)over long-term(decadal)and short-term(<10 years)scales and attempted to identify the underlying processes responsible for the observed successional patterns.Methods:The study area was the foreland of the Skaftafellsjӧkull,located in Vatnajӧkull National Park near the south coast of Iceland.We established nine transect lines at varying distances from the ice front representing surfaces of age ranging from less than one decade to over 100 years.Each transect consisted of five measurement stations of 1 m2 where we measured vegetative cover(VC),species richness(SR),and species density(SD)and calculated species evenness(SE).Measurements were made initially in 2007 and repeated at the same geographic coordinates in 2014.Results:VC increased with distance from the ice front from 16%to over 90%.SR and SD increased from the youngest pioneer community through a mid-successional stage corresponding to an age of over 60 but less than 100 years.Increased VC but declining SR,SD,and SE characterized the oldest(over 100 years)bryophyte-dominated surfaces.Species turnover,which involved forbs almost exclusively,increased moderately from early through midsuccessional sites and declined on older sites.Comparison of the measurements made in 2014 to those made in 2007 demonstrates increased SR at mid-successional sites while SD remained relatively constant.Conclusion:At a small scale,colonization is controlled by local factors such as microtopography and aspect,particularly in proximity to the glacier.At the landscape level,changes in VC and community structure are controlled by time and nutrient availability.Low nutrient levels and limited site availability favor bryophyte dominance on the oldest surfaces.The greatest community-level changes observed over the 7-year interval were increases in surface cover by mosses and low shrubs,particularly in mid-successional and older sites.These changes suggest that the community on the oldest surfaces has not yet reached equilibrium.

Intraspecific trait variation improves the detection of deterministic community assembly processes in early successional forests,but not in late successional forests

Aims Intraspecific trait variation(ITV)has been increasingly recognized to play an important role in understanding the underlying processes influencing community assembly.However,gaps remain in our understanding of how incorporating ITV will influence the relative importance of deterministic(e.g.habitat filtering,limiting similarity)and stochastic processes in driving community assembly at different successional stages.Methods We used data for eight functional traits from 55 woody species in early(24 ha)and late(25 ha)successional temperate forest plot in northeast China.We employed an approximate Bayesian com-putation approach to assess the relative contribution of stochastic processes,habitat filtering and limiting similarity in driving commu-nity structure.We then compared the results with and without intra-specific trait variation to investigate how ITV influences the inferred importance of each process.Important Findings We found that when analyzing interspecific trait variation only(i.e.without ITV),stochastic processes were observed most frequently in driving community composition,followed by habitat filtering and limiting similarity in both forests.However,ITV analyses showed that the relative importance of both deterministic processes(habitat filtering and limiting similarity)increased in early successional for-est,but remained virtually unchanged in late successional forest.Our study reveals the distinctive influence of ITV on the inference of underlying processes in a context of succession and reinforces the need to estimate ITV for making correct inferences about underlying ecological processes.

Tree phenology along a successional gradient of tropical Atlantic Forest

Aims Changes in habitat characteristics and species composition in suc-cessional gradients could determine temporal variation in phenol-ogy of second-growth forests.We evaluated phenological patterns in tree species occurring in successional forests in southern Brazil,aiming to assess community changes along succession.We tested for general patterns and phenophase seasonality of trees of forests in successional stages and for differences in occurrence,concentra-tion,frequency and duration of phenophases.Methods Vegetative(flushing and leaf-fall)and reproductive(flowering and fruiting)phenophases of 149 individuals of 29 tree species were monitored monthly,for 2 years,in a successional gradient:early-(~10 years),mid-(~30 years)and late-successional(>80 years)forests in southern Brazil.Important Findings Forests in the successional gradient exhibited shared frequency and duration of phenological phases,probably due to climatic or historical constraints in phenology.However,we found differences in the rate of occurrence and concentration of phenological phases,suggesting that habitat changes caused by succession and species turnover are additional factors affecting phenological patterns in tropical forests.

Mowing and fertilizer effects on seedling establishment in a successional old field

Aims We used a 10-year field experiment that consisted of mowing and fertilizer treatments to evaluate the role of niche limitation in seedling establishment of species from different functional groups and of varying local abundance in an old field undergoing succession.Methods Seedlings of nine different species were planted into a successional field subjected to mowing and fertilizer treatments for 10 years that resulted in different plant communities and resource availability.Species representative of the factorial combination of three functional groups(C4 grasses,C3 grasses and legumes)and three abundance categories(abundant,present,or absent in the old field)were planted in four treatments resulting from the factorial combination of annual spring mowing(mowed and unmowed)and fertilizer application(annually fertilized and unfertilized).Survivorship,relative growth rate(RGR)and biomass were measured to determine the role of niche limitation on recruitment and growth.Important Findings Mowing increased the establishment success of seedlings.Fertilization had little influence on seedling performance and survivorship.C3 grasses had the highest survivorship,while C4 grasses and legumes had equivalent RGRs,but higher than C3 grasses.By contrast,survivorship of legumes was unrelated to mowing or fertilizer,suggesting that establishment of this functional group was dependent on other,unmeasured conditions or processes.Species already present,but at low abundance,performed better than locally abundant or absent species.Propagule limitation may restrict the arrival of a species.However,recruitment and establishment was subject to niche limitation,which varied among species,functional groups and whether a species is already resident at the site and its abundance.Thus,species interactions restrict establishment during old-field succession,supporting the niche limitation hypothesis.