Rain forest expansion mediated by successional processes in vegetation thickets in the Western Ghats of India

Aim The objective of this study was to document succession from grassland thickets to rain forest, and to provide evidence for their potential as restoration tools. Location The Linganamakki region (State of Karnataka) of the Central Western Ghats of India. Method We selected thirty vegetation thickets ranging from 4 to 439 m² in area in the vicinity of rain forest. The area of each small thicket was estimated as an oval using its maximum length and its maximum width. When the shape was irregular (mostly in large thickets) the limits of the thicket were mapped and the area calculated from the map. Plant species were identified, the number of individuals was estimated and their heights measured. Results There was a progression in the thickets from early to late successional species. Small thickets were characterized by ecotone species and savanna trees such as Catunaregam dumetorum. Savanna trees served as a nucleus for thicket formation. Colonizing species were mostly bird‐dispersed. As succession proceeded in larger thickets, the proportion of evergreen, late‐successional rain forest trees increased. The species composition of the large thickets differed depending on the species composition of reproductive adults in the nearby forested areas. The species within small thickets were also found in the large thickets. The nestedness in species composition suggested that species turnover was deterministic based on thicket size. Human disturbance (leaf and wood collection by the local populations) affected the species composition and the species–area relationship of thickets. Main conclusions Vegetation thickets are nodal centres for rain forest colonization within grasslands. They expand and replace savanna. Early successional bird‐dispersed species established around savanna trees followed by late‐successional rain forest trees dispersed from the nearby forest by birds. Restoration programmes that reproduce natural successional processes such as those observed in thickets will be more successful and less expensive than the methods currently being employed, where trees are individually planted in grassland. Wood harvesting is the only factor that prevents thicket growth and coalescence and hampers forest expansion.     

Effects of competition and litter on a carnivorous plant, Drosera capillaris (Droseraceae)

Although carnivorous plants are widely recognized as being generally restricted to open habitats, tests of the effects of competition on individual performance are extremely rare. In this study, I examined the effects of the removal of herbaceous and shrub canopies on seedling density and growth, survival, and reproduction of phytometers of a small insectivorous plant, Drosera capillaris (pink sundew). I also examined the distribution of this species in relation to the occurrence of woody species in a frequently burned wet savanna in southeastern Mississippi. Killing plants and removing dead biomass increased seedling density in both open areas and shrub thickets. The removal of dead biomass following herbicide application was critical to increasing densities of seedlings. Killing plants with herbicide without also clearing residual litter and standing dead was not sufficient to increase seedling densities in shrub thickets. Although the removal of the groundcover canopy strongly influenced the density of seedlings, it had very little effect on survival, growth, and reproduction of small phytometers during a single growing season. Survival of phytometers was greater in open areas than in shrub thickets, regardless of whether the groundcover canopy was removed. Densities of both seedlings and adults were greater in open areas away from shrub thickets than beneath the woody canopies of thickets and were negatively correlated with the leaf area index of groundcover vegetation. Results of this study show that the establishment of this carnivorous plant species is limited in part by the effects of litter on seedling density in both open areas and shrub thickets.     

Tree–Shrub Interactions During Early Secondary Forest Succession in Uganda

Abstract One of the greatest challenges for ecologists this century will be restoring forests on degraded tropical lands. This restoration will require understanding complex processes that shape successional pathways, including interactions between trees and other plants. Shrub species often quickly invade disturbed tropical lands, yet little is known about whether they facilitate or inhibit subsequent tree recruitment and growth. We examined how shrubs and other vegetation (e.g., vines, grasses, herbs) affect tree recruitment, survival, and growth during the first 6 years of forest succession in Kibale National Park, Uganda. The study was undertaken in two recently logged exotic softwood plantations. We studied the successional trajectories in two recently logged areas that varied in their initial densities of trees and shrubs. Analyses suggested tree seedling presence and density were not strongly related to shrub density or height during succession. Tree sapling presence and density were positively significantly related to shrub density and height. We found little response in the tree community to experimental shrub removal, and although removal of all nontree vegetation temporarily enhanced tree growth, the effect disappeared after 2 years. Some early‐successional trees benefited from reduced competition, whereas some mid‐successional trees benefited from the presence of other vegetation. Some specific tree species responded strongly to vegetation removal. We interpret our findings in light of designing manipulations promoting forest restoration for biodiversity conservation and conclude with four tentative guidelines: (1) manage at the species level, not the community level; (2) increase facilitation for seedlings, reduce competition for saplings; (3) be cautious of assumptions about plant interactions; and (4) be adaptable and creative with new strategies when manipulations fail.     

贵州省普定县不同植被演替阶段的物种组成与群落结构特征

通过对贵州省普定县喀斯特地区不同植被演替阶段群落的调查, 研究了植被演替过程中群落物种组成和群落结构的变化。结果表明, 该地区的植被主要处于5个演替阶段, 即次生乔木林、乔灌过渡林、藤刺灌丛、稀灌草丛以及火烧干扰后的蕨类植物群落。本次调查共记录到植物365种, 隶属89科218属。其中, 蕨类植物31种, 隶属14科23属; 种子植物334种, 隶属75科195属。物种分布较多的科主要有蔷薇科、菊科、禾本科、百合科、忍冬科、唇形科、莎草科、樟科、葡萄科和水龙骨科。随着正向演替的推进, 物种丰富度增加, 群落结构趋于复杂化。藤刺灌丛与乔灌过渡林群落层次不明显, 次生乔木林分层明显。从藤刺灌丛向次生乔木林演替的过程中, 小径级个体所占比例明显降低, 高于1.3 m植物的总密度、乔木密度和藤本密度都先升高后降低, 而灌木密度呈逐渐降低的趋势。对喀斯特地区植被的恢复提出了参考措施。     

Identifying Native Vegetation for Reducing Exotic Species during the Restoration of Desert Ecosystems

There is currently much interest in restoration ecology in identifying native vegetation that can decrease the invasibility by exotic species of environments undergoing restoration. However, uncertainty remains about restoration's ability to limit exotic species, particularly in deserts where facilitative interactions between plants are prevalent. Using candidate native species for restoration in the Mojave Desert of the southwestern U.S.A., we experimentally assembled a range of plant communities from early successional forbs to late‐successional shrubs and assessed which vegetation types reduced the establishment of the priority invasive annuals Bromus rubens (red brome) and Schismus spp. (Mediterranean grass) in control and N‐enriched soils. Compared to early successional grass and shrub and late‐successional shrub communities, an early forb community best resisted invasion, reducing exotic species biomass by 88% (N added) and 97% (no N added) relative to controls (no native plants). In native species monocultures, Sphaeralcea ambigua (desert globemallow), an early successional forb, was the least invasible, reducing exotic biomass by 91%. However, the least‐invaded vegetation types did not reduce soil N or P relative to other vegetation types nor was native plant cover linked to invasibility, suggesting that other traits influenced native‐exotic species interactions. This study provides experimental field evidence that native vegetation types exist that may reduce exotic grass establishment in the Mojave Desert, and that these candidates for restoration are not necessarily late‐successional communities. More generally, results indicate the importance of careful native species selection when exotic species invasions must be constrained for restoration to be successful.     

Shrub expansion stimulates soil C and N storage along a coastal soil chronosequence

Expansion of woody vegetation in grasslands is a worldwide phenomenon with implications for C and N cycling at local, regional and global scales. Although woody encroachment is often accompanied by increased annual net primary production (ANPP) and increased inputs of litter, mesic ecosystems may become sources for C after woody encroachment because stimulation of soil CO₂ efflux releases stored soil carbon. Our objective was to determine if young, sandy soils on a barrier island became a sink for C after encroachment of the nitrogen‐fixing shrub Morella cerifera, or if associated stimulation of soil CO₂ efflux mitigated increased litterfall. We monitored variations in litterfall in shrub thickets across a chronosequence of shrub expansion and compared those data to previous measurements of ANPP in adjacent grasslands. In the final year, we quantified standing litter C and N pools in shrub thickets and soil organic matter (SOM), soil organic carbon (SOC), soil total nitrogen (TN) and soil CO₂ efflux in shrub thickets and adjacent grasslands. Heavy litterfall resulted in a dense litter layer storing an average of 809 g C m^?2 and 36 g N m^?2. Although soil CO₂ efflux was stimulated by shrub encroachment in younger soils, soil CO₂ efflux did not vary between shrub thickets and grasslands in the oldest soils and increases in CO₂ efflux in shrub thickets did not offset contributions of increased litterfall to SOC. SOC was 3.6–9.8 times higher beneath shrub thickets than in grassland soils and soil TN was 2.5–7.7 times higher under shrub thickets. Accumulation rates of soil and litter C were highest in the youngest thicket at 101 g m^?2 yr^?1 and declined with increasing thicket age. Expansion of shrubs on barrier islands, which have low levels of soil carbon and high potential for ANPP, has the potential to significantly increase ecosystem C sequestration.     

湖南大围山杜鹃灌丛的群落组成及结构特征

灌丛是陆地生态系统中最重要的自然植被类型之一, 在植被演替、生物多样性保护和维持生态平衡等方面具有重要作用。本文基于2012年的样地资料, 对湖南大围山杜鹃(Rhododendron simsii)灌丛的物种组成和群落结构进行了分析。结果表明: 群落内共有维管束植物58种, 隶属于36科50属, 其中木本植物19种, 草本植物39种, 缺乏木质藤本; 区系以温带性质为主, 兼受热带亚热带区系的强烈影响; 群落外貌矮平, 灌木层个体集中在1-2 m高度级; 植被分类上, 该群落归属于山地中生落叶阔叶灌丛, 隶属于温性落叶阔叶灌丛群系组; 生活型以地面芽和高位芽植物为主, 反映了亚热带中山山顶温凉湿润的气候特点; 群落各层次优势种明显, 具有较高的稳定性和均匀性; 高度级和基径级分析表明群落内中等大小个体具有保守的生活史策略, 优势种杜鹃为远期衰退种群。综合分析结果表明, 该杜鹃灌丛尚处于群落演替的中期或前中期, 最终将演替为亚热带中山常绿落叶阔叶混交林或常绿阔叶林。     

Enhanced nitrogen loss may explain alternative stable states in dune slack succession

Ecological theory emphasizes competitive interactions between plant species when explaining primary succession in plants. Ecosystem processes, such as nutrient accumulation, are often regarded as independent, steering successional changes without being affected by the interacting plant species. We present experimental evidence that plant species in wet dune slack systems are able to affect ecosystem processes in their favor by reducing the rate of nitrogen accumulation by coupled nitrification–denitrification, promoting their competitive position. We compared denitrification rates of two early successional species having radial oxygen loss (ROL) with two non-ROL late successional species in a mesocosm experiment. The denitrification rates were significantly higher in mesocosms planted with early successional species Littorella uniflora (P<0.001) and Schoenus nigricans (P<0.05), relative to the rates found in the presence of non-ROL species, Carex nigra and Calamagrostis epigejos . We analyzed the consequences of enhanced nitrogen loss on the competition between an early and a late successional species by means of a simple theoretical model. Our analysis revealed that early successional species capable of ROL might retard successional changes and lock the ecosystem in an unproductive state for an extended period of time. We emphasize that understanding of ecosystem processes is crucial in successful conservation of high biodiversity vegetation in wet dune slacks.     

Interactions among dwarf bamboo,litter and consumption by small vertebrates place multiple constraints on the establishment of native tree seedlings in a Japanese agricultural landscape

Aims Faunal assemblage alterations due to anthropogenic impacts have changed herbivore–vegetation relationships in various ecosystems, but the influences of small vertebrates on revegetation processes remain unclear. In a peri-urban agricultural landscape in eastern Japan that lacks large ungulates but supports small generalist herbivores, fewer native seedlings have become established in thickets dominated by native dwarf bamboo,Pleioblastus chino(Franch. et Sav.) Makino. The mechanisms limiting tree seedling establishment are unknown. Our aim here was to evaluate the influence of interactions among the dwarf bamboo, its litter cover and small vertebrate herbivores on the microsite conditions governing the establishment of native tree seedlings from different successional stages in an old field in temperate Japan.     

Shifting Mosaics in Grazed Woodlands Driven by the Alternation of Plant Facilitation and Competition

Abstract: Free-ranging large grazers, such as cattle and horses, are increasingly reintroduced to former agricultural areas in Western Europe in order to restore natural and diverse habitats. In this review we outline mechanisms by which large grazers induce and maintain structural diversity in the vegetation (mosaics of grasslands, shrub thickets and trees). This variation in vegetation structure is considered to be important for the conservation of biodiversity of various plant and animal groups. The process of spatial association with unpalatable plants (as-sociational resistance) enables palatable plants to establish in grasslands maintained by large grazers. In this way, short unattractive (thorny, low quality or toxic) species facilitate taller unattractive shrubs, which facilitate palatable trees, which in turn outshade the species that facilitated their recruitment. Established trees can, therefore, not regenerate under their own canopy, leading to cyclic patch dynamics. Since this cyclic dynamic occurs on a local scale, this contributes to shifting mosaics. The mechanisms involved in creating and maintaining the resulting shifting mosaics are described for temperate flood-plain and heathland ecosystems, including the effects on nutrient transport within grazed landscapes. How grazing leads to shifting mosaics is described in terms of plant functional types, allowing potential generalisation to other ecosystems. The resulting interaction web of grasses, unpalatable forbs and shrubs, palatable light-demanding trees and shade-tolerant trees is discussed, and was found to contain various interesting direct and indirect effects. The key process contributing to spatial diversity in vegetation structure is the alternation of positive (facilitation) interactions between plant species at one life cycle stage, and competitive displacement at another stage. Grazing thus causes directional successional sequences to change to shifting mosaics. The implications of this theory for nature conservation are discussed, including the relevant management problems, possible choices and practical solutions. We conclude that the theoretical framework outlined in this review provides helpful insights when coping with nature conservation issues in temperate woodland habitats.     

Cross-Scale Patterns in Shrub Thicket Dynamics in the Virginia Barrier Complex

Abstract To interpret broad-scale erosion and accretion patterns and the expansion and contraction of shrub thickets in response to sea level rise for a coastal barrier system, we examined the fine-scale processes of shrub recruitment and mortality within the context of the influence of ocean current and sediment transport processes on variations in island size and location. We focused on Myrica cerifera shrub thickets, the dominant woody community on most barrier islands along the coastline of the southeastern USA. Observations suggest that M. cerifera, a salt-intolerant species, is increasing in cover throughout the Virginia barrier islands, yet rising sea level in response to climate change is increasing erosion and reducing island area. Our objective was to explain this apparent paradox using pattern–process relationships across a range of scales with a focus on ocean currents and sediment transport interacting with island characteristics at intermediate scales. Multi-decadal comparisons across scales showed a complex pattern. At the scale of the entire Virginia barrier complex, modest decreases in upland area were accompanied by large increases in shrub area. Responses were more variable for individual islands, reflecting inter-island variations in erosion and accretion due to differences in sediment transport via ocean currents. Several islands underwent dramatic shrub expansion. Only for within-island responses were there similarities in the pattern of change, with a lag-phase after initial shrub colonization followed by development of linear, closed canopy thickets. Understanding the fine-scale processes of shrub seedling establishment and thicket development, in conjunction with the influence of ocean currents and sediment transport, provides a framework for interpreting island accretion and erosion patterns and subsequent effects on shrub thicket expansion or contraction across scales of time and space.     

Avian taxonomic and functional diversity in early stage of longleaf pine (Pinus palustris) stands restored at agricultural lands: variations in scale dependency

In agricultural landscapes, the longleaf pine initiative (LLPI) and the Bobwhite Quail Initiative (BQI) aim to restore longleaf pine forests and early successional habitats, respectively. The early stage of longleaf pine stands and grass and forb vegetation produced by a combination of both restoration programs (LLPI‐BQI) may form habitat conditions favorable to early successional bird species and other birds, increasing avian diversity. We investigated how the LLPI and BQI programs affected taxonomic and functional diversity of birds and abundance of early successional birds (grassland and scrub/shrub species), and what environmental characteristics were associated with the diversity and abundance of birds. Our study was performed at 41 fields in Georgia, United States, during 2001–2002 by considering environmental characteristics at two spatial scales: local‐scale vegetation features and restoration program type (LLPI or LLPI‐BQI) and landscape‐scale vegetation features and landscape heterogeneity. Functional evenness, species richness, and abundance of grassland and scrub/shrub species did not show a clear association with local‐ or landscape‐scale variables. Shannon‐Wiener diversity was slightly influenced by restoration program type (local‐scale variable) with higher value at LLPI‐BQI stands than at LLPI stands despite no significant differences in local vegetation features between those stands. Functional divergence was strongly positively associated with landscape‐scale variables. That is, niche differentiation increased with increasing shrub coverage within a landscape, reducing competition between abundant bird species and others. Our results suggest that although a combination of BQI and LLPI program may have a positive effect on avian taxonomic diversity, it is important to consider shrub vegetation cover within a landscape to improve functional diversity.     

Soil organic matter accumulation and its implications for nitrogen mineralization and plant species composition during succession in coastal dune slacks

Vegetation and soil development during succession in coastal dune slacks on Terschelling island, the Netherlands, was investigated, by comparing neighbouring ecosystems on similar substrates that had been developing for 1, 5, 35 and 76 years since the vegetation and organic soil layer had been removed. In this successional sequence, soil organic matter accumulated rapidly due to the production of litter and dead roots. N mineralization was extremely low, increasing from 0.2 g m^-2 yr^-1 after 5 years to 0.8 g m^-2 yr^-1 after 76 years. It was accompanied by a decline in the pH (KCl) in the upper 10 cm of the soil from 6.8 to 4.4. Most of the above-ground biomass accumulated in the shrub species Oxycoccus macrocarpos and Salix repens. The 5- year-old plots harboured many plant species (18 species per 0.25 m²), but plant species diversity was much lower in the older plots. It is concluded that most changes in species composition and the decline in diversity occurred because early successional plant species were gradually outshaded by the thick litter layer and the accumulated shrub biomass.     

Patterns of variability in soil properties and vegetation cover following abandonment of olive groves in Catalonia (NE Spain)

Understanding patterns and processes involved in changes in soil and vegetation after agricultural abandonment is a key issue for management policies leading to land restoration and reclamation in Mediterranean semiarid environments. We selected a number of active and abandoned fields in two regions of olive groves in Catalonia (NE Spain), in order to analyze changes in certain soil properties and vegetation cover variables, as well as their interrelationships. The soil chemical properties considered, summarized into a single PCA axis, showed significant spatial (regional) differences and no temporal (successional) pattern, indicative of the strong influence of the parent rock material. The soil physical variables examined also showed significant regional variability, but such differences could be partially explained by successional changes. The variability at the landscape level and the observed successional trends for soil physical properties are interpreted as a result of both the influence of the bedrock and contrasting management practices prior to abandonment in each region. Although we did not find significant regional or successional variability in a global measure of plant cover, we did find significant and different patterns of variability for each of the main plant functional groups considered. Regional and post-agricultural variability in soil physical properties seem to be the prime factors determining the abundance of the main plant functional groups.     

Vegetation recovery and plant facilitation in a human-disturbed lava field in a megacity: searching tools for ecosystem restoration

Unplanned urban development threatens natural ecosystems. Assessing ecosystem recovery after anthropogenic disturbances and identifying plant species that may facilitate vegetation regeneration are critical for the conservation of biodiversity and ecosystem services in urban areas. At the periphery of Mexico City, illegal human settlements produced different levels of disturbance on natural plant communities developed on a lava field near the Ajusco mountain range. We assessed natural regeneration of plant communities 20 years after the abandonment of the settlements, in sites that received low (manual harvesting of non-timber forest products), medium (removal of aboveground vegetation), and high (removal of substrate and whole vegetation) disturbance levels. We also tested the potential facilitative role played by dominant tree and shrub species. Plant diversity and vegetation biomass decreased as disturbance level increased. Sites with high disturbance level showed poor regeneration and the lowest species similarity compared to the least disturbed sites. Six dominant species (i.e., those with the highest abundance, frequency, and/or basal area) were common to all sites. Among them, three species (the tree Buddleja cordata, and two shrubs, Ageratina glabrata and Sedum oxypetalum) were identified as potential facilitators of community regeneration, because plant density and species richness were significantly higher under their canopies than at open sites. We propose that analyzing community structural traits of the successional vegetation (such as species diversity and biomass) and identifying potential facilitator species are useful steps in assessing the recovery ability of plant communities to anthropogenic disturbances, and in designing restoration strategies.     

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.     

草灌植被转变对草地生态系统及其水碳过程的影响研究进展

气候变暖和过度放牧的共同作用使全球草地出现明显的灌丛化现象,灌木去除是草地灌丛化控制的重要方式,识别这些草灌植被转变对生态系统、生态水文、土壤侵蚀和侵蚀碳流失的影响对草地可持续管理具有重要意义。综述了草地灌木入侵及其控制对植物群落和土壤功能(如土壤有机碳)的影响,以及这些草灌植被变化对生态水文、土壤侵蚀和土壤侵蚀碳流失等水碳耦合过程的影响机制。针对目前草地灌木入侵和去除对植物群落、植被格局、水土过程和功能影响研究的薄弱环节,对未来相关研究提出以下建议：(1)需深化草灌植被转变对碳、氮等生物地球化学循环的影响机制研究,(2)需重视核磁共振光谱、生物标志物、同位素等新技术和植被格局的指数与连通性等新方法在草灌植被转变的水、碳等生态效应研究中的应用,(3)需加强草灌植被格局和生态水文、土壤侵蚀与土壤侵蚀碳等水碳过程的多要素、多过程和多尺度的综合研究。本文旨在为灌丛化草地科学有效的生态恢复与多目标的土地利用管理提供理论支撑。     

Vkgetation analysis along a successional gradient from heath to oak forest

At Hjelm Hede in Denmark where an oak shrub is invading into a heathland the successional process has been studied on poor sandy outwashed plain. The study used a transect through a successional gradient from open heath to oak shrub where vegetation, major topsoil parameters and tree characteristics were analysed. The study enables us with spatial variation of key parameters to combine numerous detailed studies from the same site into an overall picture of changing vegetation and soil parameters reflecting the succession, which has been documented previously. The data were analysed by the use of Detrended Correspondence Analysis (DCA). Along the gradient from heath to forest the thick mor layer of the heathland soil decomposed, the CM ratio decreased from 25 to 15, pH increased and ammonium became available for the plants during a time span of approximately 120 years. The vegetation on the open heath was typical Danish inland heath vegetation dominated by Calluna vulgaris, Empetrum nigrum and Deschampsiaflexuosa. Under the young oak shrub the vegetation was dominated by Deschampsia Jexuosa, Trientalis europaea , and Maianthemum bijiolium. In the oldest and most forest like parts of the shrub a vegetation dominated by Holcus mollis, Anemone nemomsa and Stellaria holostea was found. Woody chamaephytes were replaced first by hemicryptophytes and later by geophytes. The vegetation's Ellenberg values indicated an increase in nitrogen availability and a decrease in acidity and light availability through secondary succession.     

Soil physicochemical properties drive the variation in soil microbial communities along a forest successional series in a degraded wetland in northeastern China

The Sanjiang Plain is the biggest freshwater wetland locating in northeastern China. Due to climate change and human activities, that wetland has degraded to a successional gradient from the original flooded wetland to dry shrub vegetation and a forest area with lower ground water level, which may result in changes in soil microbiologic structure and functions. The present study investigated the microbial diversity and community structure in relation to soil properties along that successional gradient. The soil physico‐chemical properties changed significantly with degradation stage. The Shannon diversity index of both soil bacteria (5.90–6.42) and fungi (1.7–4.19) varied significantly with successional stage (both p < .05). The community structures of soil bacteria and fungi in the early successional stages (i.e., the wetland) were significantly determined by water content, total nitrogen, and available nitrogen concentrations in soils, while those in the later successional stages (i.e., forests) were significantly structured by soil organic carbon, soil pH, and available phosphorus concentrations. These results suggest that the soil microbial structure is mainly determined by soil properties rather than by plant community such as plant species composition along successional stages.     

Keywords: ornithochory; fleshy fruit; succession; species losses; cultural biotope.

Abandonment of traditional agricultural practices in fruit orchards on hillsides in Central Germany results in successive changes in vegetation. We examined three hypotheses relating to these changes: (1) thickets of fleshy-fruited plants develop around planted trees as a result of ornithochory and local soil and site amelioration, (2) woody plants have long-term effects on soil fertility, and (3) thicket development reduces the plant-species diversity of semi-dry grassland between the trees. Field observations and nursery experiments supported the first and second hypotheses. Increased soil fertility, a seed rain of fleshy-fruited shrubs, and shrub establishment occurred mainly around planted fruit trees. Soils from old tree-planting sites remained fertile after the trees had died and disappeared. The third hypothesis was rejected because no decrease in species' richness or diversity occurred during the initial stages of thicket formation. Nevertheless, herbaceous plant species characteristic of the Festuco-Brometea community were absent from the vegetation and seed bank of shaded, nutrient-enriched sites. Conservation of semi-dry grasslands following orchard abandonment will therefore require active control of woody plants.